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FLORA 
MALESIANA 



SERIES I - SPERMATOPHYTA 

Flowering Plants 



Vol. 8, part 2 
Revisions 



INDEX TO REVISED FAMILIES 



Aceraceae 4: 3, 592 

Actinidiaceae s. str. . .4: 37 

Aizoaceae 4: 267 

Alismataceae . 5: 317; 6: 915 
Amaranthaceae 

4: 69, 593; 6: 915 
Anacardiaceae .... in press 
Ancistrocladaceae ... 4: 8 
Aponogetonaceae 4: 11; 7: 213 
Balanophoraceae .... 7: 783 

Basellaceae 5: 300 

Batidaceae 5: 414 

Betuiaceae . . 5: 207; 6: 917 

Bignoniaceae 8: 114 

Bixaceae s. str 4: 239 

Burmanniaceae . . 4: 13, 592 
Burseraceae 

5:209, 567; 6: 917; 7: 820 

Butomaceae 5: 118 

Byblidaceae 7: 135 

Callitrichaceae 4:251 

Campanulaceae . 6: 107, 928 
Cannabinaceae .... 4: 223 

Capparidaceae 6: 61 

Caprifoliaceae 

4: 175, 598; 6:928 
Cardiopteridaceae . . .7: 93 
Celastraceae 6: 227, 389, 930 
Centrolepidaceae .... 5: 421 
Ceratophyllaceae . . . .4: 41 
Chenopodiaceae 

4:99, 594; 6:932 

Clethraceae 7: 139 

Cochlospermaceae . . .4: 61 
Combretaceae 

4: 533; 5: 564; 6: 932 

Connaraceae . 5: 495; 6: 933 

Convolvulaceae . 4:388,599; 

5: 558; 6: 936; 7: 823 

Cornaceae 8: 85 

Corynocarpaceae 

4: 262; 5: 557 

Crassulaceae 4:197 

Crypteroniaceae . . . .8:187 

Cyperaceae 7: 435 

Datiscaceae 4: 382 

Dichapetalaceae 5: 305; 6: 941 
Dilleniaceae . . 4: 141; 7: 824 
Dioscoreaceae 4: 293 



Dipsacaceae 4: 290 

Droseraceae . . 4: 377; 5: 557 

Elatinaceae 4: 203 

Epacridaceae 6: 422 

Ericaceae .... 6: 469, 943 
Erythroxylaceae .... 5: 543 

Fagaceae 7: 265 

Ficoidaceae 4: 267 

Flacourtiaceae 

5: 1, 565; 6: 943; 7:827 

Flagellariaceae 4: 245 

Geraniaceae 6: 445 

Gnetaceae ... 4: 336; 6: 944 

Gonystylaceae 4: 349 

Goodeniaceae 

5: 335, 567; 6: 949; 7:827 
Haemodoraceae . . . .5:111 

Haloragaceae 7: 239 

Hamamelidaceae .... 5: 363 
Hippocrateaceae .... 6: 389 
Hydrocaryaceae .... 4: 43 
Hydrocharitaceae 

5: 381; 6: 952 
Hydrophyllaceae .... 4: 207 

Hypericaceae 8: 1 

Icacinaceae 7: 1 

Iridaceae 8: 77 

Juglandaceae 6: 143 

Juncaceae 4: 210 

Juncaginaceae 4: 57 

Labiatae in press 

Leeaceae 7: 755 

Lemnaceae 7: 219 

Lentibulariaceae . . . .8:275 
Loganiaceae ... 6: 293, 953 
Lophopyxidaceae . . .7: 89 

Malpighiaceae 5: 125 

Martyniaceae 4:216 

Molluginaceae 4: 267 

Moringaceae 4: 45 

Myoporaceae 4: 265 

Myricaceae 4: 277 

Najadaceae 6: 157 

Nyctaginaceae 6: 450 

Nyssaceae 4: 29 

Ochnaceae 7: 97 

Onagraceae 8: 98 

Oxalidaceae 7: 151 

Papaveraceae 5:114 



Passifloraceae 7: 405 

Pedaliaceae 4:216 

Pentaphragmataceae . .4:517 
Pentaphylacaceae . . .5: 121 

Philydraceae 4: 5 

Phytolaccaceae .... 4: 229 

Pittosporaceae 5: 345 

Plumbaginaceae .... 4: 107 
Podostemaceae 

4: 65; 6: 963 

Polemoniaceae 4: 195 

Pontederiaceae 4: 255 

Portulacaceae 7: 121 

Primulaceae 6: 173 

Proteaceae 5: 147 

Punicaceae 4: 226 

Restionaceae 5: 416 

Rhizophoraceae 5: 429; 6: 965 

Salicaceae 5: 107 

Salvadoraceae 4: 225 

Sarcospermaceae .... 4: 32 

Saururaceae 4: 47 

Scyphostegiaceae 

5: 297; 6: 967 
Simaroubaceae . 6: 193, 968 
Sonneratiaceae 

4:280, 513; 6: 973 

Sparganiaceae 4: 233 

Sphenocleaceae .... 4: 27 
Stackhousiaceae .... 4: 35 

Staphyleaceae 6: 49 

Stylidiaceae 4: 529 

Styracaceae 4: 49 

Symplocaceae 8: 205 

Taccaceae 7: 806 

Thymelaeaceae 

4: 349; 6: 1, 976; 7: 830 

Trapaceae 4: 43 

Trigoniaceae 4: 59 

Turneraceae 4: 235 

Typhaceae 4: 243 

Ulmaceae 8:31 

Umbelliferae . . 4:113,595; 
5:555;6:983;7:830 

Valerianaceae 4: 253 

Violaceae .... 7: 179, 831 
Xyridaceae ... 4: 366, 598 
Zygophyllaceae .... 4: 64 



LIBRARY 

THE NEW YORK BOTANICAL" GARDEN 
fifiON*. NEW XQRK 1G45.3 






^-* V 



ORB 
ULMACEAE (E. Soepadmo, Kuala Lumpur) -^ 

Monoecious or dioecious (?), evergreen, deciduous or semideciduous shrubs or 
trees, (in Mai.) unarmed and often buttressed. Growth habit (in Mai.) flush-wise, 
except for Trema and Parasponia. Indumentum of simple, bulbous-based unicellular 
hairs and/or multicellular glandular hairs. Stipules caducous or rarely rather long 
persistent and completely enclosing the bud, extrapetiolar or intrapetiolar, basally 
attached or rarely peltately attached to the nodes, free or connate. Leaves simple, 
(in Mai.) alternately arranged, petioled, pinnately nerved or triplinerved at base, 
often asymmetrical at base, entire or variously serrate. Inflorescences 1 -many- 
flowered, j, ?, 6*9. or 6*5. axillary, subterminal, or borne on leafless, older branch- 
lets or on short, lateral, leafless new shoots, paniculate, racemose, thyrsoid, cymoid, 
or rarely capitate, bracteate ; bracts minute, caducous. Flowers functionally <$, $, or 
$. — £ Flowers solitary or in condensed cymoid clusters along the rachis, sessile 
or short-pedicelled; perianth (4-)5(-7)-lobed, lobes free or variously connate, 
imbricate or induplicate-valvate in bud; stamens as many as tepals, attached to the 
base of and opposite the perianth lobes, straight or incurved in bud; anthers ovoid, 
ellipsoid or subreniform, apiculate or non-apiculate, initially tetrasporangiate, 
later becoming 2-celled, dehiscing lengthwise, introrse or extrorse; pistillode 
present or absent, if present either rather well developed or rudimentary, densely 
whitish to silvery, soft or hirsute pubescent. — $ & ~> Flowers sessile or stalked, 
solitary in the axils of the upper new leaves or arranged in various types of in- 
florescences ; perianth herbaceous or thin-coriaceous, (4-)5(-7)-lobed, lobes always 
imbricate in bud and connate at base, (in Mai.) long persistent; staminodes or 
stamens as many as perianth lobes or absent; ovary superior, 2-carpellate, (in 
Mai.) 1 -celled, sessile or stipitate; style 1, tubular, short or absent, stigmatic arms 
2, slender, often bifid to deeply lobed at the tip, adaxially papillose-stigmatic for 
their entire length; ovule 1, anatropous to hemi-anatropous, subapical, pendulous, 
bitegmic. Fruit a drupe or a samara, faintly angular or flat and winged. Seed 
mostly exalbuminous; embryo large, straight or curved; cotyledons flat-convex, 
fleshy, straight or variously folded, often foliaceous. Germination mostly epigeal. 

Distribution. There are 15 genera, c. 200 spp., widely distributed in the tropics, subtropics, 
and temperate regions of Europe (as far north as 70°, Scandinavia), Africa (South of Sahara), 
continental Asia, Malesia, Australia (Queensland and New South Wales), Pacific Islands (as far 
as Tahiti; incl. also Hawaii and the Galapagos Is.), North, Central, and South America (as far 
south as 40 \ Argentina). Fig. 1. 

Fossils. Various macrofossils (woods, drupes, and leaf-impressions) and microfossils (pollen 
grains) attributed to Ulmaceae have been reported by different authors from various localities in 
Alaska, North America, Europe, continental Asia, and Japan. Amongst the older records are 
pollen grains of the Ulmus-Zelkova-type from Golden Valley Formation in North Dakota and 
Rocky Mountains, U.S.A. (Paleocene) and wood and leaf-impressions attributed to Ulnms from 
Rocky Mountains and to Planera from Alaska (Late Paleocene). By the middle of the Eocene and 
throughout the Miocene and Pliocene fossils of Ulmaceae become more abundant (common) in 
the northern hemisphere, especially in Europe and North America. However, very little is known 
about the geological history of the family in Asia. 

Of the Malesian genera belonging to the tribe Celtoideae, fossil records of Gironniera (identifi- 
cation very doubtful) go back to Middle Eocene (Alaska), that of Cell is to Miocene (Japan), and 
Trema to Upper Oligocene (Japan). — Literature: BERRY, Tree Ancestor (1923) 146; Watari, 
Jap. J. Bot. II (1941) 385; J. I BC S^ Un. Tokyo III, 6(1952) 97; La Mottf, Mem. OeoL Soc. 
Am. 51 (1952) 112, 260, 346, 360; Pkakash & BaROHOORN, J Am. Arb. 42 (1961) 185, 347; 
GrKM ,., Tert. Angios. Hung., Ak. Kiado Budapest (1969) 83; Fkrguson, Verh. Kon. Ned. 

(31) 



32 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 1. Approximate range of Ulmaceae. 



Ak. Wet. sect. II, 60 (1971) 103; Leopold & MacGiNiTiE in Graham, Flor. & Paleofl. Asia & 
N.E. America (1972) 147; Wolfe, I.e. 200; Tanai, I.e. 235; Wolfe, Brittonia 25 (1973) 334. 

Ecology. In Malesia species of the Ulmaceae may be found in both primary and secondary 
forests, from sea-level up to 2000 m ; they are more common in the lowlands and hills. 

Apart from Ulmus lanceaefolia, Celtis rigescens, Gironniera nervosa and G. subaequalis which 
may attain up to 45 m in height and 100 cm in diameter, all species are understorey shrubs or 
trees. Of the 6 genera occurring in Malesia, species of Aphananthe, Celtis, Gironniera and Ulmus 
are basically primary forests inhabitants, though they may survive and thrive in secondary 
forests as well. Of these, Celtis and Gironniera species are the most widely spread and may be 
found growing on various types of soils, including those derived from limestone. 

In Malesia Ulmaceae are found under both everwet and more seasonal climatic conditions, 
but some show preference for one or the other. In Celtis two groups of species may be dis- 
tinguished, viz the group of C. tetrandra, timorensis and rubrovenia, and that of C. philippensis. 
The former is either semideciduous or deciduous and found mainly under a drier and more 
seasonal climate, while the latter is evergreen and grows better in more humid environmental 
conditions. Aphananthe and Ulmus species are mainly found in regions subject to a rather distinct 
seasonal climate, and they are either semideciduous or deciduous. Members of the genera 
Parasponia and Trema are pioneer plants preferring and thriving well in newly opened up 
habitats, e.g. forest clearings, thickets, roadsides, flood-plains, on volcanic ashes, etc. Fig. 10, 11, 
17. They are usually short-lived (at least in Malaya, 5-7 years) and soon will die out, particularly 
when over-grown by the other more aggressive and long-lived pioneer plants, e.g. species of 
Macaranga, Mallotus, and Grewia, etc. For this reason species of Trema and perhaps also those 
of Parasponia are usually not or rarely found in old secondary forest. It also may be noted here 
that most Trema species grow, thrive, and are more widely spread in the western parts of Malesia, 
while Parasponias are more common in the eastern parts, especially in New Guinea. 

The structure and position of the inflorescence and flowers, particularly the amount of pollen 
grains produced and the structure of the stigmas, and also the absence of nectary, seem to suggest 
that pollination is most likely affected by wind, though insects may not be ruled out altogether as 
possible agents for pollination. 

Except for Ulmus, which produces a dry, flat, winged fruit, the other Malesian genera have 
various types of fleshy drupes which turn to bright yellow, orange, or deep-red in colour when 
ripe. These drupes are most probably dispersed by various species of frugivorous birds or arbo- 
reous mammals. Alternatively, at least in some species, e.g. Celtis philippensis var. wightii which 
is very common in coastal vegetation, fruit dispersal may be carried out by water currents. In 
Ulmus the winged fruits are easily dispersed by wind. 



1977] Ulmaceae (Soepadmo) 33 

It should be noted here that there is a very high percentage of seed abortion in Malesian genera 
for reasons unknown. This is made good by the production of a great number of flowers and 
fruits, produced regularly throughout the year or at least twice a year. Except for Ulmus, the 
embryo is protected by a strong, hard and durable endocarp. In all genera endosperm is usually 
scanty or absent. 

Anatomy. For general surveys also covering the older literature see Solereder, Syst. Anat. 
Dicot. Stuttgart (1899) 860-865 and ibid. (1908) 295; Metcalfe & Chalk, Anat. Dicot. Oxford 
(1950) 1271-1278; Sweitzer, J. Arn. Arb. 52 (1971) 523-585. Additional selected references: 
Dehay, l'Appareil conducteur foliaire des Urticacees, des Moracees et des Ulmacees (Ordre des 
Urticales), Arras (1934); Janssonius, Mikr. 6, Leiden (1934) 1-308 (wood anatomy, under Urti- 
caceae); Leroy, Bull. Mus. Nat. Hist. Nat. Paris ser. 2, 18 (1946) 118-123 & 180-184 (taxonomy, 
and anatomy of Aphananthe); Den Berger, Determinatietabel Malesie, Veenman, Wageningen 
(1949) (wood identification); Rao, Govindu & Thirumalachar, J. Indian Bot. Soc. 29(1950) 
224-226 (aerial roots, Tremd) ; Janssonius, Blumea 6 (1950) 407^64 (wood anatomical affinities); 
Desch, Mai. For. Rec. 15 (1954) 618-620 (wood); Jutte, Nova Guinea n.s. 10 (1959) 241-278 
(wood); Moseley, Brittonia 25 (1973) 356-379 (anatomy and relationships). 

Sweitzer's study (I.e.) is the most up-to-date survey of leaf and wood anatomy of the Ulma- 
ceae. Although his extensive research materials included very few Malesian species his general 
conclusions are probably largely applicable to the Malesian species as well. 

The wood anatomy is indicative of the mutual affinities of all Ulmaceous genera. Shared 
characters are: predominantly simple vessel perforations, short vessel members, alternate 
intervessel pits. Fibres with simple to slightly bordered slit-like pits. Parenchyma at least partly 
vasicentric. Genera of the tribe Ulmeae (in Malesia only represented by Ulmus lanceaefolia) have 
exclusively homocellular rays. In Celteae (in Malesia all other genera) at least part of the ray 
tissue is heterocellular. 

The wood of Ulmus lanceaefolia differs from all species described in literature in lacking the 
ring porosity and the typical ulmiform arrangement of vessel clusters (original observation). 
Instead, its vessel distribution resembles the diffuse porous group of tropical Celtis species. In 
Celtis the very striking differences in vessel distribution between tropical and extratropical 
species are well documented (c/. Sweitzer, I.e.). Although from Sweitzer's and other publica- 
tions some quantitative and qualitative differences between Malesian genera of Celteae can be 
deduced, our knowledge is still based on too limited materials to allow conclusions on diagnostic 
and systematic implications. 

The leaf anatomy of Ulmaceae at the same time supports its coherence as a family and provides 
an interesting diversity, of great potential diagnostic and systematic value. All Ulmaceae share 
the dorsiventral leaf architecture. The stomata are confined to the abaxial epidermis and are of 
the anomocytic type. The indumentum includes bulbous-based unicellular trichomes the walls of 
which are usually silicified. Mineral inclusions of calcium carbonate or silica in cystoliths (with or 
without pegs) are of common occurrence. The trichome-complement, presence or absence of 
mucilage cells, crystal complement, loose or compact structure of the spongy tissue, petiole and 
midrib vasculature show a considerable diversity. Sweitzer's data and other reports from the 
literature do not yet allow a leaf anatomical characterization of the individual Malesian taxa, 
but preliminary studies are indicative that this will be possible if more material is studied. 

The entire evidence from vegetative anatomy supports the traditional placement of Ulmaceae 
in Urticales. — P. Baas. 

Palynology. Based on size, sculpturing of exine and number of pores, pollen grains of 
Ulmaceae may be divided into two major morphological types, namely the Ulmus-type and the 
( <7m-typc. In the Ulmus-lypc the pollen are oblate to subspherical, amb convex or straight; 
(4-)5(-7)-porate, 20-30 by (26~)28-38(-5l) \im, pore circular to elliptic, c. 2-3 by 3-4 [im, 
slightly thickened around its margin; exine rugulate-rcticulate. (iencra with this type of pollen 
grains are: Ampelocera, Hemiptelea, Hotoptelea, Phyllostylon, Planera, Ulmus, and Zelkova. In 

the CWm-typc the pollen is suhoblate to spherical, amb convex; (2 )3 4( 5)-poratc, pores 
circular or elliptic (elongated towards the poles), often annular and protruding, c. 2-3 by 
3 4 i/m; 17 25 by 19 30 [im; exine more or less smooth but for very line (I 1.5 |xm) scabrae. 
I his type is found in Aphununtlu\ ( rlli\, ( "hartacmc, (limnnicra, LoZtUtella, ParOSponUt, f'tcro- 



34 Flora Malesiana [ser. I, vol. 8 2 

celtis and Trema. It may be noted here that pollen grains of Gironniera, Parasponia and Trema 
are usually slightly smaller and have finer exine sculpturing than those of other genera with 
Celtis-type of pollen, while the pollen grains of the Malesian species of Trema and those of 
Parasponia parviflora Miq. are predominantly diporate. As for pollen grains, Ulmaceae are 
very closely allied to Moraceae and Urticaceae, particularly to the former. According to Nair 
(1967) the pollen type found in Ulmaceae, Moraceae and Urticaceae is derived from a tricolpate 
type of Ranalean stock. — Literature: Erdtman, Pollen Morph. & Taxon. 1 (1956) 442; Ikuse, 
Pollen Grains of Japan (1956) 62; Praglowski, Grana Palyn. 3 (1962) 45-65; Kuprianova, 
Kom. Bot. Inst. Ac. Sc. USSR 1 (1965) 54-58; Nair & Sharma, Bot. Notis. 118 (1965) 177- 
186; Straka, Pollen et Spores 8 (1966) 241-264; Nair, Rev. Palaeobot. & Palyn. 3 (1967) 
81-91; Mallik & Chaudhuri, Bull. Bot. Soc. Beng. 22 (1968) 105-108; Tsukada, Bot. Mag. 
Tokyo 81 (1968) 385-395; Rao & Lee, Pacif. Sc. 24 (1970) 255-268; Huang, Pollen Fl. PI. 
Taiwan (1972) 235; Sowunmi, Grana Palyn. 13 (1973) 145-186; Adams & Morton, Atl. 
Pollen Trees & Shrubs Canad. & U.S. 9 (1974) pi. 17; Stockmarr, Grana Palyn. 14 (1974) 
103-107; Kedves & Pardutz, Acta Biol. Szeged. 20 (1974); Hamilton, Pollen et Spores 18 
(1976) 54-57. 

Embryology. Apart from several species of Ulmus and Holoptelea very little is known about 
the sporogenesis and embryogenesis of the Ulmaceae. From a very limited information so far 
published it appears that the anthers are initially tetrasporangiate but become bisporangiate just 
before anthesis through the breakdown of the adjoining wall between the locules. The anther- 
wall development conforms with the so-called basic-type in which the parietal cells divide both 
anticlinally and periclinally to form the endothesium layer, two (Trema and Ulmus) or three to 
four {Holoptelea integrifolia) middle-layers and glandular tapetum. Simultaneous cytokinesis in 
the microspore mother-cells follows meiosis and as a result the pollen grains are initially arranged 
in either tetrahedral or decussate tetrads. At anthesis the pollen grains are either 2-celled (Holop- 
telea and Trema) or 3-celled (Ulmus). In Celtis, Holoptelea and Trema up to 80% of the pollen 
grains produced are sterile or imperfectly developed. The ovule is anatropous to hemianatropous, 
bitegmic, crassinucellar or tenuinucellar (in a few species of Ulmus) with the micropyle formed by 
both integuments (Celtis and Trema) or by the inner integument only (Holoptelea and Ulmus). In 
Holoptelea and Trema the megaspore mother-cell divides into 4 daughter cells arranged in a 
linear tetrad, and of these only the chalazal megaspore develops into Polygonum-type of embryo- 
sac. In Ulmus, however, the embryo-sac is tetrasporic and either belongs to Adoxa- or Drusa-iype 
or variation of these two types with 4-12 antipodal cells. The pollen tube enters the ovule either 
through the micropyle, the integuments or the chalaza. Endosperm formation is nuclear and 
the tissue is either diploid or triploid and later becomes cellular. Embryo development conforms 
with the Onagrad-type in Holoptelea and Solanad-type in Ulmus. Polyembryony is a common 
phenomenon, especially in Ulmus. The mature embryo is straight with broad, flat or plano- 
convex, equal or slightly unequal cotyledons in Holoptelea, Planera, Phyllostylon, Ulmus, and 
Zelkova, or curved with ascending hypocotyle and narrow, incurved or induplicate-plicate or 
variously folded cotyledons which are mostly unequal in length in Ampelocera, Aphananthe, 
Celtis, Gironniera, Parasponia, Pteroceltis and Trema. — Literature: Shattuck, Bot. Gaz. 40 
(1905) 205-223; Leliveld, Rec. Trav. Bot. Need. 32 (1935) 543-573; Capoor, Beih. Bot. 
Centralbl. 57 (1937) 233-249; Walker, Am. J. Bot. 37 (1950) 47-52; Hjelmqvist & Gazzi, Bot. 
Notis. 118 (1965) 329-360; Davis, Syst. Embryol. Angiosp. (1967) 266-267. 

Chromosomes. From various published data it seems that the chromosome number in the 
Ulmeae (Holoptelea, Ulmus, and Zelkova) is n = 14 and 2n = 28, 42, and 56, though reports of 
n = 15 and 30 have been made on Ulmus americana. In the Celtideae the number seems to be less 
constant varying from n = 10, 2n = 20, 28, 40 in Celtis (9 spp.); n = 30, 2n = 84 in Chaetacme 
(2 spp.); to n = 10, 10 + B, 18, 20, and 80 in Trema (3 spp.). It may be noted here that as for 
chromosome number, Ulmaceae seems to be closely related to Moraceae where n = 12-16, 
2n = 24, 26, 28, 42, 56, and 84, and to Urticaceae of which n = 14, 28 and 2n = 22, 24, 28, 52, 
and 84. — Literature: Krause, Ber. Deut. Bot. Ges. 48 (1930) 9-13; Planta 13 (1931) 29-84; 
Walker, Science 75 (1932) 107; Sax, J. Arn. Arb. 14 (1933) 82-84; Bowden, Am. J. Bot. 32 
(1945) 195; Darlington & Wylie, Chromos. Atlas Fl. PI. (1955) 182-183; Mangenot & 
Mangenot, Bull. Jard. Bot. Brux. 28 (1958) 315-329; Arora, Bull. Bot. Surv. India 2 (1960) 



1977] Ulmaceae (Soepadmo) 35 

305; Gajapathy, Bull. Bot. Surv. India 3 (1961) 49-51; Grudzinskaja & Zakharyeva, Bot. 
Zhurn. 52 (1967) 641-651 ; Hsu, Taiwania 13 (1967) 117-129; Mehra & Gill, Taxon 17 (1968) 
574-576; J. Arn. Arb. 55 (1974) 663-677; Fedorov (ed.), Chromos. Numb. Fl. PI. (1969) 710- 
711 ; Gadella c.s. Acta Bot. Neerl. 18 (1969) 74-83; Mehra & Hans, Taxon 18 (1969) 310-315; 
Tatayuk & Turchaninova, Tsitologia & Genetika 4 (1970) 397^01 ; Hans, Cytologia 36 (1971) 
341-345; Niehaus, Taxon 20 (1971) 355; Mehra, Nucleus 15 (1972) 64-83; Sarkar, Taxon 22 
(1973) 652. 

Chemotaxonomy. Solereder mentioned the more or less general occurrence of cystoliths 
and cystolith-like structures (Si0 2 + CaCOj) in Ulmaceae. The tendency to accumulate carbon- 
ate of lime seems to be very strong in this family; CaC0 3 is deposited in wall structures (e.g. hairs, 
cystoliths) and in cell lumina (e.g. in heartwood of Ulmus and Celtis; in seed coat cells of Celtis). 
Often oxalate of lime is also present in large amounts; solitary and clustered crystals occur in the 
family. Anatomically easily detectable internal excretion comprises also mucilage production. 
The mucilage is deposited in epidermal cells (many taxa) or in mucilage idioblasts in the meso- 
phyll of some genera and in barks and flowers of most species of Ulmus. The bark of Ulmus 
rubra Muhl. ('Slippery Elm') was used formerly as a mucilaginosum in official medicine. In 
mucilage-rich elm barks large mucilage idioblasts may develop to lysigenous mucilage cavities. 
Chemically elm bark mucilages are characterized by a high content of galacturonic acid, galac- 
tose, 3-0-methylgalactose and rhamnose. Ulmaceae are moderately strong accumulators of 
polyphenols compounds. Derivatives of caffeic acid, catechins, pro-anthocyanidins (formerly 
leucoanthocyanidins), flavonols (especially glycosides of kaempferol and quercetin) and con- 
densed (= flavanoid) tannins seem to occur more or less ubiquiteous in leaves, fruits, barks and 
woods. According to Lebreton flavonoid constituents with a trihydroxylated B-ring (in casu 
myricetin and prodelphinidin), an assumedly primitive feature, are restricted to Celtideae. 
(+)-Catechin was definitely identified in leaves, twigs and barks of European elms and its 
7-xyloside was isolated from the stem-bark of Ulmus americana L. C-Glycoflavons (tremasperin) 
occur in leaves of Trema aspera Bl., and the wood of Zelkova serrata (Thunb.) Makino contains 
large amounts of the fungistatic 6-C-glucoflavonoids keyakinin and keyakinol. Tannin contents 
of woods, barks, leaves and fruits are moderate (mostly less than 10%). There is only one report 
in literature indicating a possible co-occurrence of galli- and ellagitannins with condensed tannins 
in Ulmaceae; bark and wood of Celtis australis L. contain gallic acid and derivatives of ellagic 
acid according to Chari c.s. (1968). 

Much chemical work was performed with elm barks and especially elm woods in connection 
with 'Dutch Elm Disease'. Cadinane-type oxigenated sesquiterpenes seem to be present in the 
young wood of every species. On aging (heartwood formation) or after fungal infection, synthesis 
and accumulation of fully aromatic (cadalenal, hydroxycadalenal) and (or) o-quinonoid (the 
mansonones) cadinane derivatives take place in American elm species belonging to the sections 
Trichoptelea, Microptelea and Chaetoptelea; they seem to be absent from the sections Blepharo- 
carpus and Madocarpus in which all European elms are included. It deserves mentioning that the 
antifungal cadalenals and mansonones represent phytoalexin-like stress compounds in Ulmus, 
and occur at the same time as normal heartwood constituents in Ulmus and Zelkova (but not in 
Celtis); they are chemically identical with, or biochemically closely related to the gossypol- 
mansonone-group of constituents of many Malvaceae, Bombacaceae and Sterculiaceae (man- 
sonones were first detected in the wood of Mansonia altissima A. Chev.). It was recently shown 
that hemigossypol, the precursor of the long-known gossypol, is a phytoalexin in many malva- 
ceous plants and thatp-quinonoid derivatives of hemigossypol are engaged in the plants resistance 
against attack by several phytophagous insects (J. R. Gray c.s. J. C. S. Chem. Commun. 1976, 
109; J. A. Veech c.s. I.e. 144). As far as ecological chemistry (defensive substances) is concerned, 
Ulmaceae much resemble members of the order Malvales. Leaf, bark and wood waxes were 
investigated by several authors in recent time. They seem to consist mainly of alkanes, long-chain 
fatty acids, wax alcohols and phytosterins. Additionally pentacyclic tritcrpenes are often present ; 
^-amyrin (i), lupeol (ii), bctulin (iii), friedelin (iv), friedelanol (v), morctcnol (vi), simiarenol (vii) 
and simiarcnon (viii) were repotted from leaves and (or) barks of Celtis australis I., (iii), C. 
laevigata Wn 1 1>. (vi), Hohptelea bttegrffolia l'i anch. (iv, v), Trema gulneeruis Ficai no (reported 
as T. orientalis Bl.; vii, viii), Ulmus americana L. (ii, cstcrified with cerotinic acid) and Zelkova 



36 Flora Malesiana [ser. I, vol. 8 2 

serrata Makino (iv). The heartwood of Holoptelea integrifolia Planch, yielded 2<x-hydroxy-3- 
epioleanolic acid (G. Misra c.s. Planta Medica 27, 1975, 290); this is the only triterpenic acid 
isolated hitherto from Ulmaceae. Seeds of Ulmaceae seem to store predominantly proteins and 
fatty oils. The oils have linolic (Celtis, Chaetacme, Trema), oleic (Holoptelea) or capric (Ulmus, 
Zelkova) acids as main fatty acid. Species of Celtis and Pteroceltis accumulate small amounts of 
quebrachitol in leaves; this cyclitol could not be detected in leaves of species of Ulmus and 
Zelkova (Hemiptelea included). Alkaloid-like compounds are recorded in literature from 
members of Ampelocera, Aphananthe, Celtis, Gironniera, Trema and Ulmus, but only in the case 
of Ampelocera ruizii Klotzsch an alkaloid-like compound isolated from leaves was chemically 
identified; it proved to be an a-pyridone derivative related to trigonelline (R. H. Burnell c.s. 
Lloydia 38, 1975, 444). The foetid smell of some Celtis woods of India, Indonesia Ckaju taV) and 
Africa is caused by skatol. Several species of Ulmaceae are reported to be toxic in literature. 
Greshoff isolated a toxic bitter principle from the leaves of Aphananthe aspera (Thunb.) 
Planch. (= Homoioceltis aspera Bl.) which he compared with his streblide (from Streblus asper 
Lour.; strebloside is now known to be a cardenolide). Leaves of Trema cannabina Lour. 
(= Sponia virgata Planch.) and of T. aspera Bl. (= T. cannabina) were reported to be cyano- 
phoric; both species, however, are polymorphic with regard to cyanogenesis if the botanical 
identification of all plant samples investigated hitherto was correct. Leaves of T. aspera (= T. 
cannabina) contain another toxic principle called trematoxin; its chemical structure is not yet 
known. 

From the taxonomic point of view three facts deserve special mentioning: (1) Ulmaceae are 
generally included in Urticales; their chemistry agrees rather well with such a classification as is 
indicated by patterns of mineralisation and phenolic compounds. (2) The chemistry of Ulmaceae 
resembles members of Malvales in several respects : chemistry of stress compounds ; mucilages 
with high contents of galactose, rhamnose and galacturonic acid; some features of the poly- 
phenolic and triterpenic patterns. (3) The classification of Ulmaceae in Ulmoideae and Celti- 
doideae (Engler's Syllabus 2, 1964) or Ulmeae and Celtideae (Hutchinson, General of Flowering 
Plants 2, 1967) is not very satisfactory from the chemical point of view (see cadinane-type 
sesquiterpenes including mansonones and capric acid as main fatty acid in seed oils in Ulmus and 
Zelkova, but not in Celtis). 

For more phytochemical details and references see my 'Chemotaxonomie der Pflanzen' 6 
(1973) 545-554, 762-763, 791, 796. — R. Hegnauer. 

Taxonomy. The family name Ulmaceae was first introduced and defined by Mirbel in 1815, 
at which time it included only Celtis and Ulmus. Link (1831) proposed splitting Ulmaceae into 
two separate families, i.e. Ulmaceae to include Ulmus and related genera, and Celtidaceae com- 
prising Celtis and its allies, an opinion which was supported by Grudzinskaya (1967). However, 
all contemporary taxonomists generally agree to regard Ulmaceae as a natural taxon closely 
related to Moraceae and Urticaceae and to include these families in the order Urticales. Any 
difference of opinion is usually restricted to the inclusion or exclusion of a few genera in the 
family. In the most recent treatise, Hutchinson (1967) divided the family into two tribes, namely 
the Ulmeae (flowers bisexual, fruit not drupaceous, embryo straight, cotyledons flat or longitudi- 
nally folded) to include : Holoptelea, Planera, Phyllostylon, and Ulmus, and the Celtideae (flowers 
unisexual or sometimes bisexual, fruit drupaceous, embryo curved, cotyledons mostly variously 
folded) comprising Ampelocera, Aphananthe, Celtis, Chaetacme, Gironniera, Hemiptelea, 
Lozanella, Mirandaceltis, Parasponia, Pteroceltis, Trema and Zelkova. This subdivision was 
supported by Sweitzer (1971) who studied the anatomy of leaf and wood. However, as has been 
mentioned under Embryology and Palynology, the embryo of Zelkova is straight, and the pollen 
(also of Ampelocera and Hemiptelea) belongs to the Ulmus-type (see also Erdtman, 1956). 
Furthermore in many species of Celtis the flowers are bisexual, and in Ulmus lanceaefolia and 
U. parvifolia the flowers are either functionally male or female. This seems to indicate that the 
tribal subdivision as proposed by Hutchinson is not a clear cut case, but that Ulmaceae is a 
natural taxon. It should be noted further that the Mexican genus Mirandaceltis is in the present 
study regarded as congeneric with Aphananthe. 

As for phylogenetic relationship, there seems to be two different opinions. Bessey (1915) and 
Thorne (1968, 1973) placed Ulmaceae along with Moraceae and Urticaceae in the superorder 



1977] 



Ulmaceae (Soepadmo) 



37 




Fig. 2. Ulmus lanceaefolia Roxb. ex Wall. a. Habit, nat. size, b. fruit, • 2, c. persistent cup-shaped 
perianth, - 2, d e. embryo, nat. si/e, g. flowering twig, • 1 ,, h. cluster of * (lowers, ■ K, i. & flower, 
X 14, j. cluster of Q flowers, * 2, k I. ? flower, 6 (a /Hansi n < .s. 1 1265,^-1 Schmutz 3024,;-/ Listkr 

31). 



38 Flora Malesiana [ser. I, vol. 8 2 

Malviiflorae, and considered them as families having a very close affinity to or derived from the 
Malvales. On the other hand, authors such as Hutchinson (1967), Cronquist (1968), Takhtajan 
(1969), Sweitzer (1971), etc., are of the opinion that Ulmaceae, Moraceae, and Urticaceae are 
closely allied to or have been derived from the Hamamelidales. — Literature: Mirbel, Elem. Phys. 
Veg. Bot. (1815)905; Link, Handb. 2(1831)445; Bessey, Ann. Mo. Bot. Gard. 2 (1915) 109-164; 
Erdtman, Pollen Morph. & PI. Tax. (1956) 442^43; Grudzinskaya, Bot. Zhurn. 52 (1967) 
144-150; Hutchinson, Genera of Flowering Plants 2 (1967); Cronquist, Evol. & Class. Fl. PL 
(1968) 166-167; Thorne, Aliso 6 (1968) 57-66; Brittonia 25 (1973) 395^05; Takhtajan, Fl. PL 
Orig. & Disp. (1969) 210-212; Sweitzer, J. Arn. Arb. 52 (1971). 

Uses. 1. Timber. Throughout the north temperate regions the tough, strong and durable 
wood with attractive appearance and excellent bending quality of many species of Celtis and 
Ulmus is extensively used for various purposes including shipbuilding, panelling, furniture, boxes, 
crates, veneers, etc. and that of Zelkova and Phyllostylon for making weaver's shuttles, scales, 
piano-keys, etc. In Central America timber of Chaetoptelea (= 1 Ulmus) is used for railway 
sleepers, frames and wheels of vehicles. In Africa and India wood of Holoptelea is utilized for 
various building purposes. In Malesia and neighbouring countries except Aphananthe cuspidata, 
Celtis rigescens, C. hildebrandii, C. tetrandra, Gironniera nervosa, Ulmus lanceaefolia and a few 
others, the trees seldom reach timber size, and as a consequence very little is known about their 
usage. Of these species the timber is locally used for making planks in house-building and other 
light constructions. The soft wood of Trema and other species of Gironniera is used locally for 
making tea-chests and match-sticks, for firewood and charcoal. 

2. Bark. Due to the high content of mucilagenous substances, decoction of barks of Holop- 
telea, Parasponia, Trema and Ulmus mixed with some other ingredients is used in local folk 
medicines to cure ailments such as inflammation of mucous membrane, rheumatism, etc. The 
tough fiber is known to be used locally for making ropes. 

3. Root. Decoction of roots of Gironniera and Trema species mixed with other substances is 
used to cure sore mouth, diarrhoea, and also applied as protective medicine after child-birth. 

4. Leaves. Especially of Trema species leaves are used as fodder, though due to the presence of 
glucocides they could be poisonous if consumed in a large quantity. 

5. Fruits. In India fruits of Celtis and Holoptelea are known to be eaten. 

6. Shade trees. Trema has been used for shade in coffee and cocoa plantations in various parts 
of Asia. 

7. Soil conservation. In South Africa Trema has been planted to protect soils against erosion 
(Scheepers c.s.). As both Trema and Parasponia species come up in dense serai stands on eruptiva, 
on fresh volcanic ash, are sometimes pioneers on lavastreams, and are almost invariably an 
important constituent of thickets, serai regrowths, and secondary forest, I would emphasize that 
they may represent an untapped cheap source for soil conservation for poor, eroded soils and old 
mining lands. They have all the favourable qualities of pioneer plants, indifference to soil, pro- 
ducing abundant seed, and that already at a very early age, and furthermore they are available 
almost throughout the year. Curiously I do not know of experiments by the Indonesian Forestry 
Service in this respect. — Literature: Burkill, Diet. Econ. Prod. Mai. Pen. (1935) 513-514, 
1088-1089, 2213-2214; Metcalfe & Chalk, Anat. Dicot. (1950) 1277; Scheepers c.s. Tijd. 
Natuurwet. S. Afrika Akad. Wet. & Kunst. 8 (1968) 105-120; Sweitzer, J. Arn. Arb. 52(1971) 
525. 



KEY TO THE GENERA 

1 . Flowers always borne on bare older branches, and organized in a condensed cluster on short leafless 

lateral shoots ; perianth cup-shaped, 5-7-lobed ; ovary (fruit) stipitate. Fruit a dry, flat, winged samara. 

Embryo straight 1. Ulmus 

1. Flowers axillary, or rarely borne in a condensed capitate thyrse on older branches (Gironniera celtidi- 

folid); perianth 4-5-lobed, with the lobes free from one another except for their base; ovary sessile. 

Fruit a fleshy drupe. Embryo variously curved. 
2. Leaves triplinerved at base, or if pinnately nerved the stipules do not leave a circular scar around the 
node; lateral nerves less than 5 pairs. 

3. Stipules intrapetiolar, connate 2. Parasponia 

3. Stipules extrapetiolar, free. 



1977] Ulmaceae (Soepadmo) 39 

4. Female flowers borne in condensed, multiflowered raceme. Perianth of male flower induplicate- 

valvate. Fruit compressed, elliptic lens-shaped in cross-section 3. Trema 

4. Female flowers solitary in the axils of leaves or borne in a cymoid cluster of 2-3. Perianth lobes of 
male flowers imbricate. Fruit faintly 3-5-angular in cross-section. 
5. Male flowers borne in a 2-3-flowered cymoid inflorescence or in a much-branched paniculate, 
subterminal inflorescence. Female flowers borne in a racemose cluster of 2-10 or in a c? 5, much- 
branched racemose inflorescence; staminodes mostly present. Cotyledons broad, variously folded 

or curved 4. Celtis 

5. Male flowers organized in a condensed, multiflowered raceme. Female flowers always solitary or 
rarely borne in a 2-3-flowered racemose, o"? inflorescence; staminodes always absent. Cotyledons 

narrow, incurved 5. Aphananthe 

2. Leaves pinnately nerved ; lateral nerves more than 5 pairs ; stipules free but overlapping each other, 
in falling leaving circular scar around the node 6. Gironniera 



1. ULMUS 

Linne, Gen. PI. ed. 5 (1754) 106; Endl. Gen. PI. (1837) 276, Suppl. 2 (1842) 29; 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 259; in DC. Prod. 17 (1873) 154; Baill. Hist. 
PI. 6 (1877) 137; B. & H. Gen. PI. 3 (1880) 351 ; Hook. /. Fl. Br. Ind. 5 (1888) 480; 
Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 62; Bernard, Bull. Herb. Boiss. II, 5 
(1905) 1097; ibid. 6 (1906) 23 ; Schneider, Oest. Bot. Z. 66 (1916) 21, 65; in Sargent, 
PI. Wils. 3 (1917) 238; Gagnep. Fl. Gen. I.-C. 5 (1927) 674; Tutin, Fl. Europ. 1 
(1964) 65; Hutch. Gen. Fl. PI. 2 (1967) 147; Touw & Steen. Blumea 16 (1968) 84. 
— Fig. 2, 4-6. 

Deciduous or semideciduous trees or shrubs. Innovations densely set with greyish 
to brownish simple hairs, glabrescent. Buds ovoid-conical or obovoid-globose, 
scales imbricate, hard and tough, glabrous. Stipules extrapetiolar, caducous. Leaves 
pinnately nerved, variously serrate to crenate, thin- to thick-coriaceous and rigid, 
glabrous or variously sparsely hairy at least beneath. Flowers $ but of two kinds, 
one functionally <$ and the other functionally $, variously stalked and spirally 
arranged in fascicles of 3-15 on short lateral shoots. Perianth mostly campanulate, 
variously 4-8-lobed. Anthers glabrous, reniform, extrorse. Ovary compressed, 
sometimes stipitate; style short. Ovule 1, anatropous to amphitropous. Fruit a dry 
and compressed nutlet surrounded by a membranous reticulate-venose wing. 
Seed: endosperm absent, embryo straight with planoconvex cotyledons. Germina- 
tion epigeal. 

Distr. About 20-25 spp., distributed in Europe (as far north as 68°), W. & SW. Russia, N. & NE. 
India, Burma, China, Korea, Japan, Formosa, Indo-China, N. Thailand, and in North America from 
N. Mexico to the U.S.A. east of the Rocky Mts as far north as 60 . In Malesia: 1 sp. so far known from a 
few localities in N. Sumatra, the Lesser Sunda Is. (Flores), and Central & S. Celebes. 

As has been indicated by Schneider, I.e., there seem to be three centres of distribution, i.e. the Euro- 
pean centre (5 6 spp.), the Indian-E. Asian centre (10 15 spp.), and the North American centre (4-5 spp.). 
Fig. 3. 

Fossils. Numerous fossils (pollen grains, leaf-impressions, and wood fragments) have been reported 
from various late Cretaceous and Tertiary deposits in Europe, Russia, China, Japan, North America, and 
Greenland. Fig. 3. 

Ecol. In Malesia the genus is so far known only from areas more or less subject to a seasonal climate 
at 200 1450 m. 

Tax on. Currently there is not a single worldwide monograph of the genus available for reference. The 
latest and perhaps the most comprehensive revision since Pi aniiion's work (1X73) is that by SCHNEIDER 
(\')\f>). He distinguished 2(> spp. and recognized 5 distinct sections in the genus based on morphological 
characters derived from inflorescence, flowers, and fruits. 

When more specimens from ( hina become available for further studies, I believe the number of species 
occurring in the Indian-F. Asian centre will have to be reduced considerably. 



40 



Flora Malesiana 



[ser. I, vol. 8 J 




Fig. 3. Approximate range of Ulmus L. with number of spp. in each of the three centres, Malesian localities 

belonging to a species of the Asian centre. Fossil localities outside the present range indicated by dots ; 

adopted from Bernard, I.e.; Greguss (Tert. Angios. Hung., Ak. Kiado Budapest, 1969, 83), and La 

Motte (Mem. Geol. Soc. Am. 51, 1952, 346). 








* 









._ • t • i 



Fig. 4. Peeling bark of Ulmus lanceaefolia Roxb. ex 

Wall., x Vs (Photogr. Schmutz, 5 Nov. 1972, 

Flores, Nunang). 



1. Ulmus lanceaefolia Roxb. ex Wall. PI. As. Rar. 
2 (1831) 86, t. 200; Roxb. Fl. Ind. ed. Carey 2 
(1832) 66 {'lanci folia"); Planch. Ann. Sc. Nat. Ill 
10 (1848) 281 ; in DC. Prod. 17 (1873) 162; Kurz 
For. Fl. Burma 2 (1877) 473; Gamble, Man. Ind 
Timb. ed. 1 (1881) 342; Hook./. Fl. Br. Ind. i 
(1888) 480 ; Hemsl. J. Linn. Soc. Bot. 26 (1 894) 447 
Prain, Beng. PI. (1903) 718; Brandis, Ind. Trees 
(1906) 594; Schneider, Oest. Bot. Z. 66 (1916) 32 
in Sargent, PI. Wils. 3 (1917) 263; Merr. Contr. 
Arn. Arb. 8 (1934) 44; Touw & Steen. Blumea 16 
(1968) 84; Melville & Heybroek, Kew Bull. 26 
(1971) 24 {'lancet folia'). — U. hookeriana Planch. 
in DC. Prod. 17 (1873) 162; Engl, in E. & P. Nat. 
Pfl. Fam. 3, 1 (1888) 62. — U. tonkinensis Gagnep. 
Fl. Gen. I.-C. 5 (1927) 674. — Fig. 2, 4-6. 

Small to large tree up to 48 m, 70 cm 0, often 
with fluted trunk. Bark rough, pustulate, with large 
warty lenticels. Branchlets initially densely set with 
greyish to brownish curly simple hairs, later glab- 
rous and sparsely warty lenticellate. Buds obovoid- 
globose, c. 2-3 mm ; bracts dark brown. Stipules 
linear-lanceolate acute, c. 4-5 by 1-1 1 / 2 mm, soon 
caducous. Leaves thin- to thick-coriaceous, lanceo- 
late to ovate-lanceolate, (2-)4-6(-9) by (l-)2— 3 
(-3V2)cm (index l-l 1 ^), broadest at or slightly 
below the middle, more or less glabrous, glossy; 
base rounded to attenuate-acute, unequal; 
margin serrulate to serrulate-crenulate ; apex acute 
with blunt tip ; midrib raised beneath and flattish to 
impressed above, as the petiole initially densely 
greyish, curly hairy on both surfaces, glabrescent; 
nerves (6-)10-12(-14) pairs, subparallel, often 
rather irregularly spaced, slightly raised beneath, 
flattish to impressed above, often forked near and 
towards the leaf-margin; reticulations fine, areo- 
late; petiole (2-)3-4(-6) by 1 I 1 ~\ mm. Flowers in 
fascicles of 3-10. — Functionally J; flowers globose 
before anthesis, l'' 2 -2mm 0, subglabrous; lobes 
5-6, obovate-lanceolate, c. 2 by 1 mm; filaments 



1977] 



Ulmaceae (Soepadmo) 



41 



M J 



Uses. Very little is known about the usage of 
this species, but judging from the enormous size it 
can attain it must have been a useful timber in 
house-building, construction, etc., at least to the 
local inhabitants. 

Vern. Sumatra: pengki(h), poki, Karo-Batak, 
pongki, Toba-Batak; Lesser Sunda Is.: ngguling, 
nggulung, Flores; Celebes: mota, Bonthain. 




Fig. 5. L'lmus lanceaefolia Roxb. ex Wall, with old 

leaves at Nunang (Photogr. Schmutz, 15 Oct. 

1972, Flores). 



glabrous, slender, c. 1 mm ; anthers c. 1 by 1 I 2 mm, 
glabrous; pistillode compressed obovate-elliptic, 
glabrous. — Functionally i flowers (as seen under a 
very young fruit): perianth campanulate, lobes 
5-6, rounded-elliptic, hairy along the margin; 
filaments slender, glabrous, 3-5 mm, anthers as in 
& flower; ovary stipitate, glabrous, ± obovate- 
elliptic. Fruit obovate-elliptic, glabrous, including 
the wing 2-3 1 2 by l' 2 -2cm, stalk 5-10 mm, 
articulate, lower part hairy. 

Distr. China (?), India (E. Himalaya, Sikkim, 
Bhutan, Khasia Hills, Manipur, Assam), Bangla- 
desh, Burma (Hukong Valley, Chittagong Hills), 
Thailand (northern parts), Laos, Vietnam (Mt 
Bavi); in Malesia: N. Sumatra (Gajo- & Karo- 
Batak Lands), Lesser Sunda Is. (Flores), and Cele- 
bes (Poso; Bonthain). Fig. 7. 

Ecol. Scattered tree in lowland to submontane 
forest, 200- 1450 m. In Thailand it is confined to 
forests along streams and in Flores it has been 
found on limestone. Fl. fr. in the northern hemis- 
phere Febr. April; in Flores Nov. 

Tax on. V. lanceaefolia is very closely allied to 
U. parvifolia JaOQ. from China and Japan. It differs 
from the latter by its narrow leaves with a shorter 
petiole, serrulate to serrulatc-crcnulatc margin, 
and fewer lateral nerves, obovoid-globosc buds, 
the campanulate perianth of the functionally V 
flower, and the reticulate venation of the fruit; see 
fig. 2. 



Fig. 6. Ulmus lanceaefolia Roxb. ex Wall., 

leafless, in flower, ± x 2 / 3 , at Nunang (Photogr. 

Schmutz, 5 Nov. 1972, Flores). 




Fig. 7. Range of Ulmus lanceaefolia Roxb. ex Wall. 



42 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 8. Parasponia rigida Merr. & Perry, a. Habit, x 2 / 3 , b. twig-tip with stipules, c. connate stipues 
from inside, d. detail of leaf undersurface, all x 3, e. 6 flower, f. ditto m LS g young ? flower^ mature 
? flower i. ditto in LS, all x 12, j. fruit, k. ditto in LS, both x 12 (a-f ANU 6463, g-k Schodde 4828). 



1977] 



Ulmaceae (Soepadmo) 



43 



2. PARASPONIA 

Miq. PI. Jungh. (1851) 68; Fl. Ind. Bat. 1, 2 (1859) 218; Bl. Mus. Bot. 2 (1856) 65; 
Planch, in DC. Prod. 17 (1873) 194; Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 
65; J. J. Smith in K. & V. Bijdr. 12 (1910) 662; Back. & Bakh./. Fl. Java 2 (1965) 
12; Hutch. Gen. Fl. PI. 2 (1967) 149. — Fig. 8, 10-11. 

Shrubs to medium-sized trees. Bark grey-brown, smooth to finely fissured; inner 
bark fibrous and tough. Innovations with dense appressed, silvery to greyish hairs. 
Stipules intrapetiolar, connate into a bifurcate unit and together enclosing the 
terminal bud, caducous. Leaves (in Mai.) triplinerved at base, concolorous, above 
non-scabrous to variously scabrous, mostly glabrous except for the midrib and 
lateral nerves, lower surface variously pubescent. Inflorescences axillary, 6*, 9, or 
6*$, much-branched, many-flowered, paniculate or thyrsoid, including the bracts 
densely short greyish appressed-pubescent. Flowers 5-merous. — 6* Flower ± 
globose, perianth lobes imbricate in bud; stamens glabrous, introrse; filaments 
subulate, glabrous ; anthers reniform to subglobose, sub-basifixed, glabrous ; pistil- 
lode obovoid-conical, compressed, surrounded by hirsute hairs at its base. — $ 
Flower ovoid-conical; staminodes absent; ovary ovoid, slightly compressed; stig- 
matic arms short, simple; ovule anatropous. Drupe ovoid, slightly compressed 
pericarp fleshy and fibrous, endocarp hard and stony. Seed: endosperm scanty or 
copious; embryo curved, cotyledons equal, hypocotyle ascending. 

Distr. 5 spp., in Polynesia (Tahiti) and Melanesia (Fiji, New Hebrides, Solomons); in Malesia: New 
Guinea (incl. New Britain), Moluccas (Ternate, Banda), Philippines, Celebes, Lesser Sunda Is. (Lombok, 
Bali), Java, and S. Sumatra (Palembang). Fig. 9, 12. 




Fig. 9. Approximate range of the genus Parasponia Miq. (line); localities of P. andersonii (Planch.) 
Planch, (dots) and P. melastomatifolia J. J. S. (triangles). 



Ecol. In New Guinea and the Pacific islands the genus is found as a pioneer plant invading and occupy- 
ing newly available habitats from the lowland up to 2000 m. In Java the same situation was described by 
F. W. ( j a.V)n from natural regeneration on volcanic ash of Mt Kelud, together with Trtma (Mull. Jard. 
Hot. Ht/g III, 13, 1935, 509). fig. 10, II. Recorded U a pioneer on lavastrcams of Mt Hatur in Bali by 
Dt VOOOD (Tirop. NatUUT 29, 1940, 48, f. 12). Grows well in all ty|H.-s of soils. Including volcanic ash and 
limestone, very often gregariously together with Trema spp. and forming a dense thicket on ridges, hills 
and along river-banks. 



44 



Flora Malesiana 



[ser. I, vol. 8 2 



Taxon. Parasponia is morphologically very similar to Trema but can easily be distinguished from the 
latter by its imbricate perianth lobes of the male flowers and intrapetiolar, connate stipules enclosing the 
terminal bud. 

KEY TO THE SPECIES 

1. Basal nerves running up throughout the length of the leaf or nearly so. 

2. Leaf thick-coriaceous, lower surface densely set with soft erect hairs; margin distinctly serrate; 
reticulations prominent beneath. Inflorescence $ or ?, at anthesis condensed and shorter than the 
petiole 1. P. rigida 

2. Leaf chartaceous to thin-coriaceous, lower surface glabrous or sparsely appressed-hairy ; margin 
finely serrulate to subentire; reticulations obscure. Inflorescence $ or <J$, at anthesis lax and longer 
than the petiole 2. P. melastomatifoiia 

1. Basal nerves running up to x \ 2 - 1 \ i the length of the leaf. 

3. Leaf thick-coriaceous, upper surface strongly rugose and scabrous, lower surface densely pubescent; 
midrib, nerves, and reticulations prominent beneath 3. P. rugosa 

3. Leaf chartaceous to thin-coriaceous, upper surface not or hardly rugose nor scabrous, lower surface 
sparsely appressed pubescent or glabrous; midrib, nerves, and reticulations only slightly raised 
beneath. 
4. Leaf elliptic-lanceolate, more or less glabrous; nerves more than 4 on each side, straight and ascend- 
ing at a narrow angle (less than 40°) from the midrib. Inflorescences mostly <J$, very rarely <$ or $ 

4. P. parvifiora 
4. Leaves ovate to ovate-elliptic, underneath sparsely appressed pubescent; nerves less than 4, usually 3 

on each side, arcuating at a wider angle (c. 45-60) from the midrib. Inflorescences <$ or $, rarely c?2 

5. P. andersonii 



1. Parasponia rigida Merr. & Perry, J. Arn. Arb. 
22 (1941) 254. — Fig. 8. 

Small tree, up to 10 m, 10 cm 0. Branchlets 
initially densely silver-appressed-hairy, glabrescent 
and sparsely warty lenticellate. Stipules ovate- 
lanceolate, 8-10 by 2-3 mm. Leaves elliptic to 
ovate-lanceolate, thick-coriaceous, (5-)8-10(-12) 
by {\ l ! 2 -)2-VI 2 {-A l ! 2 ) cm (index 2 1 / 2 -3 1 /2), broadest 
at or below the middle ; base rounded to subcordate, 
mostly symmetrical ; margin serrate ; apex acute to 
acuminate; above more or less glabrous, rugulose 
and scabrous, beneath densely soft-hairy; midrib 
and nerves strongly raised beneath, flattish to im- 
pressed above; reticulations subscalariform, dis- 
tinct beneath; petiole 10-12 by 2-3 mm, terete. 
Inflorescences 3 or ?, very rarely <??, 10-50-flowered, 
at anthesis condensed, c. l l 2 -l cm long, usually 
shorter than the petiole. — $ Flowers 1-2 mm 0, 
perianth lobes narrow-ovate, concave, c. 2 by 1 mm, 
appressed-pubescent outside; filaments c. 1 mm; 
anthers ellipsoid, c. 1 by x \ 2 mm, pistillode obovoid- 
conical, c. 1 by l l 2 mm. — ? Flower ovoid, c. 2 by 
1 mm; perianth lobes ovate-acute, c. 3 / 4 by l j 2 mm, 
sparsely pubescent outside; ovary c. 2 by 1 mm, 
stigmatic arms spreading, long-papillose, c. x / 2 - 
1 mm. Drupe ovoid-globose, c. 3-4 mm 0, turning 
orange to red when ripe. Endosperm copious. 

Distr. Malesia: New Guinea. Fig. 12. 

Ecol. Dominant pioneer tree in secondary 
vegetation on ridges, also in mossy forest and on 
limestone hills, 1000-2000 m. Fl.fr. Jan.-Dec. 

Vern. Besukan, Hattam lang., golan, Finsch- 
hafen dial., Morobe Distr. 

2. Parasponia melastomatifoiia J. J. Smith, Nova 
Guinea 8, 2 (1914) 891, t. 158. — P. simulans Merr. 
& Perry, J. Arn. Arb. 22 (1941) 255. 

Shrub to small tree, up to 6 m, with spreading 
and brittle branches. Branchlets initially densely 
appressed-silvery-hairy, glabrescent and sparsely 
lenticellate. Stipules ovate-elliptic, 4-5 by 2-3 mm. 
Leaves elliptic-lanceolate to ovate-elliptic, (5-)6-8 



(-10) by (VI 2 -)2'l 2 -Z'l 2 {-A'l 2 )cm (index 2-3), 
broadest at or slightly below the middle; charta- 
ceous to thin-coriaceous, above more or less 
glabrous, hardly scabrous and often with mineral 
deposits on the epidermis, beneath sparsely ap- 
pressed-hairy especially on midrib and nerves, or 
completely glabrous; base more or less rounded, 
symmetrical; margin finely serrulate to ± entire; 
apex acute; midrib and nerves slightly raised 
beneath, flattish to impressed above; nerves 1-2 
pairs, the basal ones running throughout the length 
of the leaf or nearly so, at an acute angle of less than 
45 J ; reticulations subscalariform, inconspicuous 
on both surfaces; petiole (5-)8-12(-15) by 1 mm, 
sulcate, sparsely appressed-pubescent. Inflores- 
cences 3 or 6*?, paniculate, 10-20-flowered, at 
anthesis lax, 2-3 cm long and across, axes c. 1 mm 
0, densely appressed-pubescent; bracts ovate, 
c. IV2 by 1 mm, sparsely appressed-pubescent 
outside. — <$ Flowers c. 1-2 mm 0, perianth lobes 
ovate-acute, c. V2-I by 1 j 2 mm, sparsely appressed- 
hairy outside; filaments c. 1 mm, anther sub- 
globose, c. 1 by V2 mrn > pistillode obovoid, c. 1 / 2 -l 
by V2 mm. — $ Flower ovoid, c. 1-2 by 1 mm; 
perianth lobes ovate-acute, c. 1 / 2 -l by 1 U- 1 ( 2 mm, 
sparsely appressed-hairy outside ; ovary ovoid, c. 2 
by 1 mm ; stigmatic arms c. 1 mm, incurved. Drupe 
ovoid-globose, c. 3 by 2 mm. Endosperm scanty. 

Distr. Malesia: New Guinea. Fig. 9. 

Ecol. Common and dominant in serai vegetation 
in gullies and river-banks, 200-1000 m. Fl. fr. 
Jan.-Dec. 

Vern. Kwatoro, Onjob lang., Northern Distr., 
E. New Guinea. 

3. Parasponia rugosa Bl. Mus. Bot. 2 (1856) 66. — 
P. aspera Bl. I.e. 66; Miq. Fl. Ind. Bat. 1, 2 (1859) 
218. — Trema vulcanica Merr. Philip. J. Sc. 7 
(1912) Bot. 260. — Trema philippinensis Elmer, 
Leafl. Philip. Bot. 9 (1934) 3218. — P. parviflora 
(non Miq.) Steen. Philip. J. Sc. 91 (1962) 507. — 
Fig. 10-11. 



1977] 



Ulmaceae (Soepadmo) 



45 



Shrub or medium-sized tree, up to 20 m, 30 cm 
0, with spreading branches. Bark smooth, grey- 
brown; inner bark tough, brownish. Branchlets 
densely, silvery, appressed, long-hairy, subglab- 
rescent. Stipules ovate-lanceolate, (6-)8(-10) by 
2-3 mm. Leaves ovate-lanceolate, (7— )8— 10(— 12) 
by (2-)2 1 2 -3' 2 (-5) cm (index 2Vj-3), broadest at 
or below the middle, thick-coriaceous; above 
rugose and scabrous, sparsely hairy especially on 
midrib and nerves, beneath densely silvery hairy by 
soft, erect hairs; base rounded to cordate, equal to 
slightly unequal; margin serrate, apex acute to 
acuminate, the acumen up to 2 cm; midrib and 
nerves strongly raised and prominent beneath, 
flattish to impressed above; nerves 2-4 pairs, 
arcuate and ascending, the basal ones running up to 
l li- 2 U tne length of the leaf; reticulations dense, 
areolate, prominent beneath; petiole 7-10 by 
2-3 mm, subterete, densely appressed-pubescent. 
Inflorescences $ or 3?, 10-50-fiowered, densely sil- 
very appressed-hairy, at anthesis condensed, axes 
c. V2-IV2 cm long, 1-2 mm ; bracts ovate-acute, 
c. IV2-I mm. — c? Flower c. 1-2 mm 0; perianth 
lobes ovate-acute, c. 1-2 by 1 mm, sparsely hairy 
outside; filaments l-l\' ? mm, anthers ovoid-reni- 
form, c. 1 mm ; pistillode ovoid, compressed, 
c. 1-2 by 1 mm. — $ Flower ovoid-conical, c. 1-1 V2 
by 1 mm; perianth lobes narrow ovate-acute, c. 1- 
P/2 by 1 mm, sparsely appressed-hairy outside; 
ovary ovoid, c. 1-2 by 1 mm, stigmatic arms c. 
1 mm, spreading. Drupe ovoid, 2-3 by 2 mm, 
turning red when ripe. Endosperm copious. 

Distr. Malesia: East Java (Mts Kelud & 
Lamongan), Lesser Sunda Is. (Bali, Lombok), 
Philippines (Luzon, Leyte, Mindanao), Celebes 
(near Makassar; Tondano, Menado), Moluccas 
(Ternate, Banda), New Guinea (W. & E. High- 
lands and Morobe Distr., incl. New Britain). 
Fig. 12. 




Fig. 10. Pioneer vegetation on the volcanic ash of 
Mt Kelud, East Java, of Saccharum spontaneum 
and Parasponia rugosa Bl. (Photogr. Clason). 



Ecol. Rather common and often dominant or 
co-dominant pioneer plant in serai vegetation on 
various types of soils including volcanic ash, 50- 
1900 m. Fl.fr. Jan.-Dec. Fig. 10, 11. 

Uses. Strips of the inner bark are used as ropes 
in house and fence building by local inhabitants. 

Vern. Java: anggring, anggris, J; Philippines: 
analdung, If.; Moluccas: kayu kuli, Banda; New 
Guinea: wanep, Enga lang., W. Highlands Distr., 
la karabi, W. Nakanai, New Britain. 




Fig. 1 1. Older pioneer forest on Mt Kelud of Parasponia rugosa Bl., Trema, Cyathea contaminans, and 

Amomum (Photogr. Clason). 



46 



Flora Malesiana 



[ser. I, vol. 8 2 



4. Parasponia parviflora Miq. PI. Jungh. (1851) 
69; Fl. Ind. Bat. 1, 2 (1859) 218, t. 16; Bl. Mus. 
Bot. 2 (1856) 65, f. 35; Planch, in DC. Prod. 17 
(1873) 194; J. J. Smith in K. & V. Bijdr. 12 (1910) 
663, p.p excl. syn. P. aspera Bl.; Back. & Bakh. 
/. Fl. Java 2 (1965) 12. — P. similis Bl. Mus. Bot. 2 
(1856) 66. 

Small to medium-sized tree, up to 15 m. Branch- 
lets initially densely silvery or grey appressed-hairy, 
glabrescent, smooth. Stipules ovate, 5-10 by 
2-4 mm, sparsely hairy outside. Leaves lanceolate 
to narrow ovate-lanceolate, (3-)5-8(-10) by 
(l-)2-3(-3V2) cm (index 3-4), broadest at or below 
the middle ; chartaceous to thin-coriaceous, above 
± glabrous, not scabrous, beneath initially 
appressed-hairy, later glabrous except for the 
midrib and nerves; base rounded, more or less 
equal; margin finely serrate, apex acute; midrib 
and nerves slightly raised beneath, impressed and 
inconspicuous above; nerves 4-6 pairs, straight, 
ascending and parallel, at a narrow angle (30-40°), 
basal ones running up to ± half the length of the 
leaf; reticulations fine, subscalariform, indistinct 
on both surfaces ; petiole terete, densely appressed- 
hairy, 5-10 by 1 mm. Inflorescences 6\ 9, or 6*9, 
5-30-flowered, at anthesis condensed, shorter than 
or as long as the petiole, as the bracts densely 
short-hairy; bracts ovate-acute, I-IV2 by 1 l 2 -l mm. 
— 6* Flowers glabrous, 1-2 mm ; perianth lobes 
c. I-IV2 by 1 /2 mrr K filaments l / 2 -l mm, anthers 
subglobular, c. 1 by V2 mm ; pistillode ovoid- 
conical, compressed, 1-1 V2 by 1 I 2 mm. — 9 
Flowers ovoid-conical, c. 2 by P/2 mm » ± glabrous 
except for the inner base of the perianth lobes; 
perianth lobes ovate-acute, c. 1 by 1 I 2 mm; ovary 
ovoid, c. IV2 by 1 mm; stigmatic arms spreading, 
c. V2-I mm. Drupe ovoid-conical, slightly com- 
pressed, IV2-2 by IV2 mm. Endosperm scanty. 




Fig. 12. Localities of Parasponia rugosa Bl. (dots), 

P. parviflora Miq. (triangles), and P. rigida Merr. 

& Perry (squares). 

Distr. Malesia: S. Sumatra (Palembang, very 
rare), Java (common). Fig. 12. 



Ecol. In secondary or serai vegetation on ex- 
posed habitats, also in teak forest, often rather 
common and dominant locally especially on soils 
derived from volcanic ash, 500-2000 m. Fl. fr. 
Jan. -Dec. 

Vern. Kurai, k. lelaki, k. tjangkreng, S, anggring, 
anggris, anggrung, J. 

5. Parasponia andersonii (Planch.) Planch, in 
DC. Prod. 17 (1873) 193. — Sponia andersonii 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 336; See- 
mann, Fl. Vit. (1867) 235; Parham, PI. Fiji Isl. 
(1972) 133. — P. paucinervia Merr. & Perry, J. 
Arn. Arb. 20 (1939) 324. 

Shrub to medium-sized tree, up to 15 m and 
30 cm 0. Branches spreading and drooping, 
initially densely set with erect but soft, silvery hairs, 
subglabrescent and sparsely warty lenticellate. 
Bark smooth to nodular, grey-brown; inner bark 
fibrous, tough, orange to brownish. Stipules ovate- 
acute, sparsely hairy outside, 6-10 by 3-4 mm. 
Leaves ovate to elliptic, thin-coriaceous, (5— )8-12 
(-14) by (2-)3-4(-6) cm (index 2-3), broadest 
below or at the middle; above subglabrous, scab- 
rous, often covered with mineral deposits, beneath 
sparsely set (rarely rather densely) with short and 
soft hairs especially on midrib and nerves; base 
rounded to subcordate, equal, rarely unequal; 
margin serrate, apex acute to acuminate; midrib 
and nerves slightly raised beneath and impressed 
above; nerves 3-4 pairs, arcuating and ascending 
at an angle of 45-60°, basal extending up to c. 2 / 3 
the length of the leaf; reticulations fine, subscalari- 
form, rather distinct below; petiole (7-)10-15(-20) 
by 1-2 mm, densely set with silvery, soft, erect 
hairs, flat or sulcate. Inflorescences 6\ 9, or rarely 
69, 10-30-flowered, at anthesis condensed or lax, 
shorter than or as long as the petiole, including the 
bracts densely silvery, soft-hairy; bracts ovate- 
acute, c. 1 by V2 mm. — 6* Flowers c. \ l l 2 -2 mm ; 
perianth lobes ovate-elliptic, c. 1 by 1 / 2 mm ; 
stamens glabrous; filaments c. 1 mm, anthers 
subreniform to subglobular, c. 1 by x \ 2 mm ; 
pistillode subovoid-conical, c. IV2 by 1 I 2 mm. — 9 
Flowers ovoid-ellipsoid, c. \ l j 2 by 1 mm; perianth 
lobes ovate-acute, c. 1 by '/ 2 mm; ovary ovoid, 
slightly compressed, c. 1 by '^mm; stigmatic 
arms c. 1 I 2 mm, spreading and short-papillose. 
Drupe ovoid, slightly compressed, 2-4 by 2-3 mm. 
Endosperm copious. 

Distr. Polynesia (Tahiti), Melanesia (Fiji, New 
Hebrides, Solomons, very common), ? New 
Caledonia (no specimen seen but cf. Guillaumin, 
Fl. Nouv.-Caled. 1948, 94); in Malesia: New 
Guinea (several islands off Madang and Milne 
Bay) and New Britain. Fig. 9. 

Ecol. Primary as well as secondary forests, on 
various types of soils including limestone, 
0-1500 m. Fl.fr. Jan.-Dec. 

Uses. In the Solomons the bark is reputed to 
have medicinal properties. 

Vern. New Britain: ip, ivu; Solomons: bulasisi, 
bulsisi, Kwara lang. ; Fiji : ndroi, ndrou. 



1977] Ulmaceae (Soepadmo) 47 

3. TREMA 

Lour. Fl. Coch. 2 (1790) 562; Bl. Mus. Bot. 2 (1856) 58; Bth. Fl. Austr. 6 (1873) 
157; B. & H. Gen. PI. 3 (1880) 355; Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 65; 
Bernard, Bull. Herb. Boiss. II, 6 (1906) 31, maps 19-21 ; J. J. Smith in K. & V. 
Bijdr. 12 (1910) 649; Rendle, Fl. Trop. Afr. 6, 2 (1917) 10; De Wit, Bull. Bot. 
Gard. Btzg III, 18 (1949) 184; Hutch. Gen. Fl. PI. 2 (1967) 148; Elias, J. Am. Arb. 
51 (1970) 37, f. 2; Soepadmo in Whitmore, Tree Fl. Mai. 2 (1973) 420. — Sponia 
Commers. ex Lamk, Diet. 4 (1795) 138; Endl. Gen. PI. 4 (1837) 276; Planch. 
Ann. Sc. Nat. Ill, 10 (1848) 264; in DC. Prod. 17 (1873) 195. — Fig. 13, 16-17. 

Trees or shrubs, often buttressed and with spreading and drooping branches, 
monoecious. Innovations variously and densely set with simple bulbose-based hairs 
or/and with short multicellular capitate-glandular hairs. Terminal buds ovoid- 
conical, enclosed by overlapping but free extrapetiolar, caducous stipules. Leaves 
penninerved ; above ± glabrous, variously scabrate, beneath glabrous, subglabrous, 
or variously densely set with bulbous-based hairs and/or with short multicellular 
glandular hairs; base triplinerved, cordate to acute, often unequal-sided; margin 
variously serrate or denticulate; apex acute to acuminate or caudate; petiole sul- 
cate. Inflorescence axillary, paniculate or thyrsoid, many-flowered, condensed or 
lax at anthesis, ^, $, 6*$> densely and variously pubescent; bracts minute, ovate- 
acute, caducous. — <$ Flower globular; perianth 4-5-lobed, lobes induplicate- 
valvate in bud, boat-shaped; stamens glabrous, introrse; filament subulate, glab- 
rous, incurved in bud; anthers subglobular to reniform, glabrous, dorsifixed near 
the base; pistillode present, hirsute at base. — $ Flower ovoid; perianth 4-5-lobed; 
staminodes absent or very rarely present; ovary ovoid, (in Mai.) glabrous, slightly 
compressed, sessile ; style short ; ovule ana- to amphitropous. Drupe ovoid or sub- 
globose, (in Mai.) slightly compressed, glabrous; exocarp fleshy and fibrous, 
endocarp stony and very hard. Seed with a rather scanty or copious endosperm ; 
embryo curved or nearly involute; hypocotyle ascending; cotyledons equal. 
Germination epigeal. 

Distr. About 10-15 spp., widely distributed throughout the tropics and subtropics. In Asia (with 6-7 
spp.) from the warmer parts of the Himalayas, extending north-eastwards to China (incl. Hainan, Hong- 
kong, Formosa) and S. Japan and south and south-eastwards through India, Burma, Thailand, Indo- 
China, and Malesia to the tropical and subtropical parts of Australia and the Pacific islands as far east as 
Tahiti (31 N-37 S). In Africa (with 3-4 spp.) it occurs south of the Sahara to S. Africa and Madagascar 
(22 N-28 S). In America (with 4-5 spp.) the genus is known from Central & S. Florida and Mexico, 
extending south-eastwards through Central America, Bermuda, and the Bahamas, the Greater Antilles 
and southwards to South America as far south as the northern parts of Argentina (26° N-25° S). In 
Malesia: 4 spp., widely spread. Fig. 14, 15. 

Ecol. Throughout its range of distribution the genus seems to grow well and often gregariously in 
newly opened up habitats on various types of soils ranging from heavy laterite to limestone soils and soils 
derived from volcanic ash (fig. 17), from sea-level up to 2000 m. 

Pollination is probably affected by wind and small insects. 

The fruits which turn orange, red or black when ripe are dispersed by various species of bulbuls. 

HON (Bull. Jard. Bot. Btzg III, 13, 1935, 509, f. Ill) reported that following the frequent 
eruptions of Mt Kclud, Trema spp. together with Parasponia spp. formed a dominant association in the 
regrowths on volcanic ash. 

Tax on. The genus is homogeneous and closely related to Parasponia and Celtis. This is corroborated 
by the anatomy of the wood and leaves. Reports on the cytology are, however, suggesting that the number 
of chromosomes is not constant. 

I.mhryology. Very little is known about the sporogencsis and embryogencsis of the genus. A preliminary 
study carried out recently on Ircma lannahina and I . tomentosa in the Malay Peninsula indicates that the 
development of the anther and microspores follow the so-called dicotyledon-type, and that of the embryo- 
sac conforms with the Polygonum-lypc. 



48 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 13. Trema orientalis (L.) Bl. a. Habit, with functionally $ flowers, x 2 / 3 , b-e. 6* flowers at various 

stages of development, all x 8, f. 5 inflorescence, x 2 / 3 , g. $ flower, h. older $ flower, 2 tepals removed, 

I. ditto in LS, j-k. mature fruit, 1. ditto, exocarp halfway removed, showing stone, m. ditto in LS, n. embryo, 

o. detail of lower leaf surface, all x 8 (a-e BW 13889,/-/ BW 7019, j-n Brass 6496). 



1977] 



Ulmaceae (Soepadmo) 



49 




Fig. 14. Approximate range of Trema Lour. 




Fig. 1 5. Species density of Trema Lour, in Indo-Australia ; above the hyphen the number of endemic spp. 

below it the number of non-endemic spp. 



Chromosomes. A few counts on the chromosome number which have been reported by various cytolo- 
gists suggest that cytogenetically the genus is rather variable. In Trema poUtoria from India n 10 B 
(MtHRA & Gill, Taxon 17, 1968, 574; J. Am. Arb. 55, 1974, 663); in T orientals n = 18 (Arora, Bull. 
Surv. India 2, I960, 305), or n = 20 (OAJAPATHY, ibid. 3, 1961,49; Hsu, Taiwania 13, 1967, 117), or 
n ■ 10 (Mmi'A <t HANS, Taxon IK, 1969, 310; Hans, Cytologia 36, 1971, 341); and in T. tomentosa 
(cited as T. amboinensb) D = 10 or 80 (Hans, /.<•.; Mihka, Nucleus 15, 1972,64). 

S/'rv (fie delimitation has proved to he dillicult and has led to more than 50 names in the genus. This was 
partly due to the various interpretations of the eaily described species. I here is still no unanimity of 
opinion about the number of good species in the continents. In Africa, for example. ENGLER (Pfl, Welt Afr. 
}(1), 1913, ihcstiiK lumber foi Africa at 5 7, following Blum (1856), but Rbndlb (1917) and 

hiimii (Kcw Hull. 19, 1964, 143) accept only one, either under the specific name /'. gutntttwii or T. 
orientals. 



50 Flora Malesiana [ser. I, vol. 8 2 

In absence of a critical, reliable world monograph there is a similar uncertainty about the number of 
species in the neotropics and in Indo-Malesia. For Malesia out of 20-25 published names of species and 
varieties, only 4 spp. are recognized here. 

The proliferation of name giving in Malesia is mainly due to the fact that Trema spp. have a growth 
habit of continuously producing lateral and terminal new shoots on which flowers and fruits are borne. 
Many specimens collected were from these young shoots in which the indumentum and leaf-shape is often 
different from that of mature leaves. For accurate identification leaves, inflorescences, and fruits of mature 
specimens are essential. Besides, the indumentum was in earlier descriptions mostly derived from low 
magnification observations, but to differentiate sterile material of T. orientalis and T. tomentosa the 
difference in the indumentum becomes only clear under at least 40 x magnification. It is impossible to 
name young sterile specimens. 

KEY TO THE SPECIES 

{Based on mature leaves, inflorescences, and fruits) 

1. Petiole (7-)10-15(-25) mm. Leaves broad ovate-elliptic, rarely narrow lanceolate (but then either 
glabrous or silvery brown tomentose beneath), (3— )5— 10(— 1 3) by (l l l z -)2-4(-5 l / 2 ) cm (and then glab- 
rous or sparsely pubescent beneath) or (5-)10-15(-19) by (2-)3-7(-10) cm. Inflorescence d, ?, or c?$, 
at anthesis lax, 10-100-flowered, 1 x / 2 -5 cm long. Mature fruit 3-5 by 2-4 mm. 
2. Leaves (3-)5-10(-13) by (l 1 / 2 -)2-4(-5 1 / 2 ) cm, glabrous or rarely sparsely pubescent beneath, char- 
taceous to thin-coriaceous; base rounded, rounded-attenuate, truncate, or very rarely subcordate, 
mostly symmetrical; nerves 2-4(-5) pairs. Inflorescence always with a slender axis. Mature fruit 

c. 3 by 2 mm, orange or red in colour 1. T. cannabina 

2. Leaves (5-)10-15(-19) by (2-)3-7(-10) cm, densely and variously hairy beneath, thin- to thick- 
coriaceous; base cordate, subcordate or rounded, mostly asymmetrical; nerves 4-8 pairs. Inflores- 
cence with a stout or slender axis. Mature fruit black, 3-5 by 3-4 mm. 
3. Leaves beneath (fig. 16) completely covered with matted glaucous to silvery straight appressed or 
curly, erect hairs and short, multicellular capitate-glandular hairs (epidermis invisible even under 
high magnification), very often distinctly discolorous with the upper surface darker in colour and 

weakly scabrate. $ Inflorescence up to 2 l j 2 cm. Fruit ± globular 2. T. orientalis 

3. Leaves beneath (fig. 16) densely or sparsely (but not completely) set with velvety greyish-brown erect 
hairs only (epidermis clearly visible between the hairs even under low magnification), mostly con- 
colorous, dark-brown to blackish in dried specimens; upper surface strongly scabrate. <$ Inflores- 
cence up to 5 cm. Fruits ovoid, compressed 3. T. tomentosa 

1. Petiole (2-)3-6 (-8) mm. Leaves narrow ovate-lanceolate, (4-)5-6(-ll) by (lV 4 -)2-3(-4) cm, beneath 
densely tomentose by short, matted, rufous, 1 -celled and multicellular glandular hairs (hoary). 
Inflorescence c? or c?9, at anthesis condensed and shorter or as long as petiole, 5-15-flowered. Mature 
fruit subglobose, 2-3 by 2 mm 4. T. angustifolia 

1. Trema cannabina Lour. Fl. Coch. 2 (1790) 563; Laut. et var. pallida (Bl.) Laut.; Hand.-Mazz. 

Merr. Trans. Am. Phil. Soc. 24, 2 (1935) 131 ; De Symb. Sin. 7 (1929) 107. — T. virgata (Planch.) 

Wit, Bull. Bot. Gard. Btzg III, 18 (1949) 184; Bl. Mus. Bot. 2 (1856) 59; J. J. Smith in K. & V. 

Back. & Bakh./. Fl. Java 2 (1965) 12; Soepadmo Bijdr. 12 (1910) 652; Laut. Bot. Jahrb. 50 (1913) 

inWhitmore, Tree Fl. Mai. 2(1973)421. — Celtis 313, incl. var. pubigera (Bl.) Laut. I.e. 315; 

amboinensis Willd. Sp. PI. 4,2(1805)997. — Celtis Schneider in Sargent, PI. Wils. 3 (1917) 289; 

commersonii Brongn. in Duperrey, Voy. Bot. Coq. Ridl. Fl. Mai. Pen. 3 (1924) 319 ; Gagnep. Fl. Gen. 

Phan. (1829) 215. — Sponia commersonii (Brongn.) I.-C. 5 (1927) 686; Hand.-Mazz. Symb. Sin. 7 

Decne, Nouv. Ann. Mus. Hist. Nat. Ill, 3 (1834) (1929) 106; Corner, Ways. Trees (1940) 694; Li, 

498; Planch. Ann. Sc. Nat. Ill, 10 (1848) 317; in Woody Fl. Taiwan (1963) 109. — T. amboinensis 

DC. Prod. 17 (1873) 198. — Sponia timorensis (Willd.) Bl. Mus. Bot. 2 (1856) 61, quoad nomen, 

Decne, Nouv. Ann. Mus. Hist. Nat. Ill, 3 (1834) excl. syn. et sched.; Merr. Int. Rumph. (1917) 187. 

498; Planch. Ann. Sc. Nat. Ill, 10 (1848) 318; in — T. viridis (Planch.) Bl. Mus. Bot. 2 (1856) 58. 

DC. Prod. 17 (1873) 196; Miq. Fl. Ind. Bat. 1,2 — T. glabrescens (Planch.) Bl. I.e. 58. — T. cari- 

(1859) 216. — Sponia amboinensis (Willd.) nata Bl. I.e. 59. — T. pallida Bl. I.e. 60. — T. 

Decne, Nouv. Ann. Mus. Hist. Nat. Ill, 3 (1834) pubigera Bl. I.e. 60. — T. morifolia Bl. I.e. 59; 

498, quoad nomen. — Sponia virgata Planch. Ann. Laut. Bot. Jahrb. 50 (1913) 318. — Sponia pallida 

Sc. Nat. Ill, 10 (1848) 316; in DC. Prod. 17 (1873) (Bl.) Miq. Fl. Ind. Bat. 1, 2 (1859) 215; Planch. 

195, incl. var. major Planch. I.e. 196; Miq. Fl. in DC. Prod. 17 (1873) 196. — Sponia carinata 

Ind. Bat. 1, 2 (1859) 216; Kurz, For. Fl. Burma 2 (Bl.) Miq. Fl. Ind. Bat. 1, 2 (1859) 215; Planch, in 

(1 877) 469. — Sponia glabrescens Planch. Ann. Sc. DC. Prod. 1 7 (1 873) 202. — Sponia pubigera (Bl.) 

Nat. Ill, 10 (1848) 317; Miq. Fl. Ind. Bat. 1, 2 Miq. Fl. Ind. Bat. 1, 2 (1859) 216; Planch, in DC. 

(1859) 217. — Sponia viridis Planch. Ann. Sc. Prod. 17 (1873) 197. — Sponia morifolia (Bl.) 

Nat.III,10(1848)319. — T. commersonii (Brongn.) Planch, in DC. Prod. 17 (1873) 196. — Sponia 

Bl. Mus. Bot. 2 (1856) 60. — T. timorensis (Decne) vieillardii Planch. I.e. 201. — Sponia aspera var. 

Bl. I.e. 60; Hook. /. Fl. Br. Ind. 5 (1888) 483; viridis (Planch.) Bth. Fl. Austr. 6 (1873) 158. — 

Hemsl. J. Linn. Soc. Bot. 26 (1894) 452; Laut. T. orientalis var. amboinensis (Willd.) Kurz, For. 

Bot. Jahrb. 50 (1913) 317, incl. var. carinata (Bl.) Fl. Burma 2 (1877) 469, quoad nomen. — T. vieil- 



1977] 



Ulmaceae (Soepadmo) 



51 



lardii (Planch.) Schltr, Bot. Jahrb. 36 (1905) 
31. 

Shrub or small much-branched tree up to 6 m, 
15 cm 0. Bark smooth, grey-brown. Branchlets 
slender, spreading, often drooping, initially densely 
silvery-hairy, glabrescent and sparsely lenticellate. 
Stipules linear-lanceolate, 5-7 by 1-2 mm. Leaves 
chartaceous to thin-coriaceous, narrow ovate- 
caudate to broad ovate-acute, or elliptic-lanceolate, 
(3-)5-10(-13) by (l 1 2 -)2-4(-5 1 2 ) cm (index 2-3 
(-4)), broadest below or at the middle; base 
rounded to attenuate and acute, rarely subcordate, 
slightly contracted and more or less symmetrical; 
margin serrulate to denticulate for its entire length ; 
apex with a sharp tip ; above glabrous and variously 
scabrate, beneath glabrous or sparsely appressed- 
hairy ; midrib and nerves raised beneath, impressed 
above; nerves (2-)3-4(-5) pairs, arcuate and sub- 
parallel, basal ones running up to ± */a the length 
of the leaf; reticulations fine, subscalariform, 
obscure to visible beneath; petiole (5— )8— 12(— 15) 
by 1-2 mm, glabrescent. Inflorescence <$ or <£?, 
with slender axes, 10-15-flowered, at anthesis lax, 
c. 1— 2 1 2 cm long, densely greyish appressed-hairy ; 
bracts ovate-acute, c. 2-3 by 1 mm. — $ Flowers 
c. 1-2 mm 0, outside sparsely hairy, glabrescent; 
perianth lobes 4-5, membranous, oblong-lanceo- 
late, c. 1—1 x 2 by ' 2 -l mm; filaments c. 1 mm, an- 
thers c . 1 by l / a mm ; pistillode obovoid, compressed, 
c. 1 by l /j mm - — ? Flowers c. lV 2 -2 by 1-1 l / 2 mm; 
perianth lobes mostly 5, membranous, glabrous, 
ovate-acute, c . 1-1 l /j by l / a mm ; staminode absent ; 
ovary c. 1 by l /a mrr >; stigmatic arms spreading or 
incurved. Drupe 2-3 by 2 mm, turning deep-orange 
or red when ripe. Endosperm copious. 

Distr. Burma, China, Formosa, Hainan, Indo- 
China, Thailand, common throughout Malesia to 
Australia, Melanesia (Solomons, New Caledonia, 
New Hebrides), W. Polynesia (Fiji, Samoa), and 
Micronesia. 

Ecol. Common as a pioneer in newly opened up 
habitats along roadsides, edges of forests, re- 
growths, thickets, and in young secondary vegeta- 
tion, from sea-level up to 1200 m. Fl.fr. Jan. -Dec. 
At least in Malaya pollination is affected by wind 
and by small insects (diptera). Ripe fruits are dis- 
persed by various species of bulbuls. 

Taxon. In Malesia there seem to be three rather 
but not completely distinct entities. These can be 
defined as follows: (i) specimens which have a 
completely glabrous, chartaceous and narrow- 
ovate leaf of (3- )5-8(- 10) by (l' 2 -)2-3(-4) cm with 
an index of 2 1 2 3, more or less non-scabrate upper 
surface, and 2-3 lateral nerves (T. cannabina and 
T. virgata); (ii) specimens with a thin-coriaceous, 
narrow ovate-lanceolate to elliptic-lanceolate leaf 
of (6-)7-10(-12) by (l l ; r )2-3(-3 1 / 2 )cm, with an 
index of 4-5, slightly scabrate upper surface and 
sparsely hairy lower surface, and 4-5 lateral nerves 
which are straight and ascending and forming a 
narrow angle (less than 40 ) with the midrib (T. 
timorensis, T. virgata var. scabra, and T. cannabina 
var. scabra), (iii ) those with a broad ovate and coria- 
ceous leaf of (H )') IK 13) by (3 )4 4 1 2 ( 5' ,) cm, 
with an index of 2" 2 - 3. rugose and slightly scab- 
rate upper surface and sparsely pubescent beneath, 
and 3 4 lateral nerves forming a broad angle (more 
than 45 ) with the midrib (7. gla!" I trktis, 

and T. vieillardii). Various intermediates arc how- 



ever present, making it difficult to recognize them 
as distinct infra-specific taxa. 

Vern. Malaya: minirong, mingkirai, M; S. 
Sumatra: dilung, M, Palembang; Java: anggrung, 
J ; N. Borneo : bintanong, Murud ; Lesser Sunda Is. : 
ridong kue, Flores, pipaka, Alor; Moluccas: 
loli-sawu, Halmaheira; Solomons: bulasisi, Kwara. 

2. Trema orientalis (L.) Bl. Mus. Bot. 2 (1856) 62; 
Bth. Fl. Austr. 6(1873) 158; Hook./. Fl. Br. Ind. 5 
(1888) 484; J. J. Smith in K. & V. Bijdr. 12 (1910) 
655, p.p., excl. syn. T. commersonii et T. griffithii; 
Laut. Bot. Jahrb. 50 (1913) 320, p.p., incl. var. 
rigida (Bl.) Laut. I.e. 322, excl. var. viridis et var. 
amboinensis; Corner, Ways. Trees (1940) 694, 
pi. 211, p.p. excl. syn.; de Wit, Bull. Bot. Gard. 
Btzg III, 18 (1949) 189, p.p., incl. var. bicornis De 
Wit, I.e. 190, excl. var. bicolor et syn. T. angusti- 
folia, T. imbricata et T. velutina; Back. & Bakh./ 
Fl. Java 2 (1965) 12, p.p., excl. syn. T. amboinensis 
auct. non (Willd.) Bl.; Soepadmo in Whitmore, 
Tree Fl. Mai. 2 (1973) 421. — Celtis orientalis 
Linne, Sp. Pi. 2 (1753) 1044; Roxb. Fl. Ind. ed. 
Carey 2 (1832) 65. — Celtis rigida Bl. Bijdr. (1825) 
486. — Celtis discolor Brongn. in Duperrey, Bot. 
Voy. Coq. Phan. (1829) 215, pi. 47B. — Sponia 
discolor (Brongn.) Decne, Nouv. Ann. Mus. Hist. 
Nat. Ill, 3 (1834) 498; Planch. Ann. Sc. Nat. Ill, 
10 (1848) 324; in DC. Prod. 17 (1873) 201. — 
Sponia orientalis (L.) Decne, Nouv. Ann. Mus. 
Hist. Nat. Ill, 3 (1834)498; Planch. Ann. Sc. Nat. 
Ill, 10 (1848) 323; in DC. Prod. 17 (1873) 200. — 
Sponia rigida (Bl.) Decne, Nouv. Ann. Mus. Hist. 
Nat. Ill, 3 (1834) 498; Planch. Ann. Sc. Nat. Ill, 
10 (1848) 336; Miq. Fl. Ind. Bat. 1, 2 (1859) 217. — 
Sponia argentea Planch. Ann. Sc. Nat. Ill, 10 
(1848) 323; in DC. Prod. 17 (1873) 201. — Sponia 
wightii Planch. Ann. Sc. Nat. Ill, 10 (1848) 322; 
Wight, Ic. 6 (1853) t. 1971. — T. argentea 
(Planch.) Bl. Mus. Bot. 2 (1865) 58. — T. bur- 
mannii Bl. I.e. 62. — T. rigida (Bl.) Bl. I.e. 61. — 
T. scaberrima Bl. I.e. 63. — T. wightii (Planch.) 
•l. I.e. 58. — T. discolor (Brongn.) Bl. I.e. 58; 
Laut. Bot. Jahrb. 50 (1913) 319. — Sponia scaber- 
rima (Bl.) Miq. Fl. Ind. Bat. 1, 2 (1859) 217; 
Planch, in DC. Prod. 17 (1873) 202. — Sponia 
burmannii (Bl.) Planch, in DC. 17 (1873) 200. — 
Fig. 13, 16. 

Shrub to large tree, 3-36 m, 10-90 cm 0. But- 
tresses, if present, up to l'^m. Bark smooth to 
finely fissured, lenticellate, grey-brown or whitish- 
grey. Branchlets, stipules, petioles, and inflores- 
cences densely set with appressed and matted or 
erect silvery to glaucous 1 -celled hairs and short 
multicellular glandular hairs. Stipules linear- 
lanceolate to ovate-acute, 3-4 by 1-2 mm. Leaves 
thin- to thick-coriaceous, often rigid and brittle, 
ovate, ovate-lanceolate to narrow elliptic, lan- 
ceolate, (6-)10-15(-18) by (lV 2 -)2'/ 2 -6(-10) cm, 
index (2-)3^4(-5'/ 2 ), broadest at or mostly below 
the middle, mostly discolorous, above dull grey- 
brown or grey-green in dried specimens, scabrate 
and sparsely set with bulbous-based hairs, beneath 
densely tomentose by a combination of silvery, 
glaucous or grey-brown, appressed 1 -celled hairs 
and shorter multicellular glandular hairs (fig. 16); 
base cordate, rounded, or sometimes truncate, 
often contracted, asymmetrical or symmetrical; 
margin serrate to denticulate for its entire length; 



52 



Flora Malesiana 



[ser. I, vol. 8 2 



apex acute to acuminate-caudate; midrib and 
nerves raised beneath and impressed above; nerves 
4-6(-8) pairs, the lowest pair arcuating and run- 
ning up to l l 2 - 2 l 3 the length of the leaf; reticulations 
subscalariform to subareolate, sometimes strongly 
raised and distinct beneath; petiole (7— )10— 15(— 1 8) 
by 1-2 mm, densely short pubescent. Inflorescences 
either <$ or ? borne on separate vegetative branches, 
a much-branched panicle or thyrse, at anthesis 
lax or condensed, axes 1-2 mm thick ; bracts ovate- 
acute, 2-3 by 1 mm. Flowers 5-merous. — c? 
Inflorescences up to 3-5 cm long, 20-100-flowered; 
cJ flower c. lV2-2mm 0; perianth lobes ciliate, 
IV2-2 by 1mm; filaments l-l'/jmm, anthers 
c. 1 by 7a mm > pistillode obovoid-conical, com- 
pressed, 1-1 V2 by V2-I mm - — $ Inflorescences 
5-15-flowered, l 1 /2-2 1 /2cm long, axes 1-2 mm 
thick; 9 flower c. 2-3 by 1-2 mm; perianth lobes 
ovate acute, c. 1— 1 1 / 2 by V2 mm > ciliate and densely 
short pubescent, glabrescent; staminode absent; 
ovary ovoid-conical, c. 2 by 1 mm ; stigmatic arms 
slender, c. \-\ l l 2 mm., spreading. Drupe 3-5 by 
2-4mm, turning black when ripe. Endosperm 
scanty to copious. 

Distr. TTropical Africa, SE. Asia (Ceylon, 
India: from W. Himalayas to Bombay and Mala- 
bar; Burma, Thailand, Indo-China, China, also 
Hainan, Formosa, to S. Japan), through Malesia 
to Queensland, Melanesia (Solomons), Micronesia 
(Marianas), and Polynesia (Fiji, Tonga, Tahiti). 
In Malesia: Malay Peninsula and Sumatra (rather 
rare), Java (rather common in the hills and sub- 
montane regions), Lesser Sunda Islands (rare), 
Borneo (common), Philippines (rather common), 
Celebes (rather rare), Moluccas (rare), and New 
Guinea (incl. New Britain, rare). 

Ecol. In W. Malesia and continental Asia the 
species is more common in the hills and montane 
regions between 600-2000 m, whereas in E. 
Malesia, Australia, and Pacific Islands it is more 
common in the lowlands. The ripe fruits which 
turn to deep purple or black are dispersed by. 



various species of birds, particularly bulbuls. 
Fl. fr. Jan.-Dec. 

Taxon. Three rather but not completely distinct 
entities may be recognized. These are: (i) specimens 
from continental Asia and W. Malesia which have 
been variously identified as T. orientalis, rigida, 
argentea, and wightii by previous authors. They 
are characterized by: thick-coriaceous, broadly 
ovate to ovate-elliptic leaves with grey-brown to 
glaucous indumentum, slightly asymmetrical to 
symmetrical cordate, subcordate or rounded base, 
rugose upper surface, and acute to acuminate 
apex ; and by a relatively larger fruit of c. 4-5 by 
3-4 mm and stouter inflorescence axes. 

(ii) Specimens from S. Japan, Formosa, Hainan, 
the Philippines, New Guinea, Micronesia, Mela- 
nesia, and Polynesia, and Australia, which have 
been included in the so-called T. discolor, charac- 
terized by: thin-coriaceous, narrow-ovate leaves 
with strongly asymmetrical cordate base, hardly 
scabrate upper surface, short and matted silvery to 
grey-brown indumentum, lax inflorescence with 
slender axes, and fruits c. 3-4 by 2-3 mm. 

(iii) A few specimens from scattered localities in 
S. China, Thailand, Sumatra, and Borneo, which 
have been described by De Wit (1949) as T. orien- 
talis ssp. bicornis, characterized by: very narrow, 
ovate-lanceolate thin-coriaceous leaves with silvery 
appressed and matted dense indumentum on the 
lower surface and non-scabrate upper surface, 
6-8 pairs of nerves, and the shorter and few- 
flowered inflorescence. 

Several intermediates are present however, 
making formal infraspecific distinction not ad- 
visable. 

Vern. Sumatra: endrung, ndirung, Karo, 
indarung, Pajakumbu, bandorung, Tapanuli, lando- 
jung, Simelungun, endelung, Palembang, Bencoo- 
len, magelong, nelung, Bencoolen, neriung, Lam- 
pong. Java: gorai, kuraj, S, anggrung, njampu, J. 
Lesser Sunda Is.: lenggung, Bali, redong, Flores, 
tabelah, W. Sumbawa. Borneo: randagong, 




Fig. 16. Indument of lower leaf-surface, strongly enlarged. Left: only unicellular hairs of Trema tomentosa 

(Roxb.) Hara, between which the epidermis is visible. Right: T. orientalis (L.) Bl., with long unicellular 

hairs and crowded multicellular crisped hairs covering the epidermis. 



1977] 



Ulmaceae (Soepadmo) 



53 



tandago, Dusun; bingkirai, E. Kutai. Philippines: 
anadgong, Bis. Celebes: ngawoi, Malili, mawa, 
Bonthain, kantu, Toradja, tajapu. Moluccas: rufu, 
Ternate, laei, Tidore, soka soka, E. Ceram. West 
New Guinea: bisuwai, Hattam lang., karara, 
Ambai, kaniem, mier, Kebar lang. 

3. Trenla tomentosa (Roxb.) Hara, Fl. E. Himal. 
2 (1971) 19; Soepadmo in Whitmore, Tree Fl. Mai. 
2 (1973) 423. — Celtis orientalis {non L.) Bl. Bijdr. 
(1825) 485. — Celtis amboinensis {non Willd.) 
Brongn. in Duperrey, Bot. Voy. Coq. Phan. 
(1829) 212, pi. 47 A, p.p., excl. specim. ex Ventenat, 
Amboina. — Celtis tomentosa Roxb. Fl. Ind. ed. 
Carey 2 (1832) 66. — Sponia amboinensis (Willd.) 
Decne, Nouv. Ann. Mus. Hist. Nat. Ill, 3 (1834) 
498, quoad specim.; Planch. Ann. Sc. Nat. Ill, 10 
(1848) 321; Miq. Fl. Ind. Bat. 1, 2 (1859) 216; 
Planch, in DC. Prod. 17 (1873) 198. — Celtis lima 
(non Sw.) Blanco, Fl. Filip.2 (1837) 139. — Sponia 
griffithii Planch. Ann. Sc. Nat. Ill, 10 (1848) 324. 
— Sponia tomentosa (Roxb.) Planch. I.e. 336. — 
Sponia velutina Planch. I.e. 327, p.p., excl. specim. 
Cuming 1232 ex Luzon. — Sponia blancoi Planch. 
I.e. 327; Miq. Fl. Ind. Bat. 1, 2 (1859) 218. — T. 
griffithii (Planch.) Bl. Mus. Bot. 2 (1856) 58. — 
T. blancoi (Planch.) Bl. I.e. 58. — T. imbricata 
Bl. I.e. 63. — T. velutina (Planch.) Bl. I.e. 58; 
Gagnep. Fl. Gen. I.-C. 5 (1927) 689 ; Li, Woody Fl. 
Taiwan (1963) 109. — T. amboinensis (Willd.) 
Bl. Mus. Bot. 2 (1856) 61, quoad specim.; Bth. Fl. 
Austr. 6 (1873) 159; Hook./. Fl. Br. Ind. 5 (1888) 
484; K. Sch. & Laut. Fl. Schutzgeb. (1900) 264; 
J. J. Smith in K. & V. Bijdr. 12 (1910) 659, p.p., 
excl. syn. Celtis amboinensis Willd. et Trema 
burmannii Bl.; Merr. En. Born. (1921) 217; 
Ridl. Fl. Mai. Pen. 3 (1924) 319. — Sponia 
imbricata (Bl). Planch, in DC. Prod. 17 (1873) 
199. — T. orientalis var. amboinensis (Willd.) 
Kurz, For. Fl. Burma 2 (1877) 469, quoad specim.; 
Laut. Bot. Jahrb. 50 (1913) 321. — T. orientalis 
{non (L.) Bl.) Merr. Sp. Blanc. (1918) 121. — T. 
dielsiana Hand.-Mazz. Symb. Sin. 7 (1929) 106; 
P'ei, Bot. Bull. Ac. Sin. 1 (1947) 289. — Fig. 16. 

Shrub to medium-sized tree of 5-15(-24)m, 
5-3O(-50) cm 0. Bark grey-brown, smooth to 
finely fissured, lenticellate. Branchlets, inflores- 
cences, petioles, stipules, and underside of leaves 
densely and thickly set with greyish, erect, velvety 
hairs. Stipules linear-lanceolate, c. 5 by 1 mm. 
Leaves thin- to thick-coriaceous, broadly ovate to 
ovate-elliptic, (5-)8-15(-19) by (2-)4-7(-9) cm, 
index 2' 2 -3, broadest mostly below the middle; 
more or less concolorous, drying dark-chocolate 
brown to blackish brown ; above strongly scabrate; 
base cordate, rarely subcordate or rounded, mostly 
strongly asymmetrical, rarely symmetrical; margin 
serrate throughout, apex acute to acuminate- 
caudate, acumen sharp, 1-3 cm; midrib and nerves 
raised beneath (often very strongly), impressed and 
hairy above; nerves 4-6 pairs, ascending and 
subparallel, at an angle of • 45 , the lowest pair 
running to the length of the leaf; reticu- 

lations subscalariform to suhatcolate, often rather 
distinct beneath; petiole I— l*/ a cm by I 2mm, 
densely pubescent, Infloresa-nns .either 

on the same or on different vegetative branches; 
bracts ovate-acute, r. I by '/j mm. - - At anthesis 
(J and J j axes of the inflorescences lax, 2*/j 4'/j cm 



long, 20-100-flowered; c? flower c. lV2-2mm 0; 
perianth lobes mostly 5, elliptic, c. l x / 2 by 1 mm; 
filaments c. 1 mm, flat, glabrous, anthers c. 1 by 
'/2 mm; pistillode obovoid-ellipsoid, compressed, 
IV2 by V2 mm - — ? Inflorescence 1-2 cm long, axes 
1-2 mm thick, 5-15-flowered; 9 flower c. 2 by 
1 mm; perianth lobes 4-5, ovate-acute, c. 1 by 
\ 2 mm; staminode mostly absent, if present 
strongly reduced in size and non-functional; ovary 
c. l'/2 by 1 2~1 mm, stigmatic arms slender, c. 1 mm, 
spreading. Drupe c. 3 by 2 mm, maturing black. 
Endosperm copious. 

Distr. East tropical Africa, Madagascar, SE. 
Asia: Pakistan, India, Bangladesh, Burma, Thai- 
land, Indo-China, China (incl. Hainan), Hong- 
kong, Formosa, Ryu Kyu Is. (Okinawa), through- 
out Malesia to Queensland, Melanesia (New 
Caledonia), Micronesia, and Polynesia (Fiji, 
Tonga, and Hawaii). 

Ecol. Common in the lowlands and hills, at sea- 
level up to 1000 m, as a pioneer plant invading and 
occupying newly opened up habitats on all kind of 
soils, including limestones. Fl. fr. Jan. -Dec. At 
least in Malaya pollination is affected by wind and 
small insects (diptera). The ripe black fruit is dis- 
persed by various species of birds. 

Tax on. Evidently, T. tomentosa is closely allied 
to T. orientalis, and it is possible that, when more 
field data become available in the future, the former 
may prove to be only representing a juvenile 
ontogenetical form of the latter. Except for a few 
specimens from the Philippines {e.g. Whitford 
681, BS 37313, 48355, Elmer 8417) and from New 
Guinea {e.g. ANU 2075, 2752, 6240, Hartley 
10937, Manner & Street 270, NGF 29353, 
Schodde 1419, and BW 16510) in which the leaves 
are thick-coriaceous and with a more or less sym- 
metrical base and pale grey-brown in colour, 
specimens of T. tomentosa can be easily dis- 
tinguished from those of T. orientalis by the charac- 
ters mentioned in the key. Fig. 16. It is also interest- 
ing to note that according to Hans (Cytologia 36, 
1971, 341) and Mehra (Nucleus 15, 1972, 64) the 
chromosome number in T. tomentosa is n = 10 or 
80, whereas that of T. orientalis is n = 10, 18, or 20. 

Vern. Malay Peninsula: minarong, m$ndarong, 
mgngkirai, M. Sumatra: bingkirai, Gajo, endelung, 
Palembang, e2maha, Enggano, hana{w)e, Batak, 
kamisen silai, Simalur, mangkirai, Pajakumbu, 
manghirei, minkirei, Lingga, intuitu sabu, Djambi, 
{n)d$r(r)ung , Karo-Batak, randurung, Toba, sang- 
kiraja, Batak, tindjau, Riouw. Java: anggrung, J, 
kuraj, k. awewena, S. Lesser Sunda Is.: riaong, 
damot, Flores, ruka parak, Sumba. Borneo: Sara- 
wak : murieng, Bidajuh, kirininog, [ban, tuku baroh, 
Land Dayak; N. Borneo: anjalakat, Kedayan; 
Brunei: balek balik angin jantan, balik origin, 
rundagong, Brunei, btntonong, Murut, damai, 
Suluk, enttmon, I ban, llndagong, Kedayan, lunda- 
gong, Dusun Tambato & Kayan, lundagong, salt' 
muak, Dusun, randagong, Tenggara, rinaagong, 
Dusun Labuk; E. Borneo; bangirai, bangkiral, 
tjalundung, 1 . Kutai; w. Borneo: tngkirai, hum. 
Philippines: anaginong, Mang., anugdon, Tag-Bis., 
anabtong, hanagdong, Tag., Icarangyan, karayang- 
yang, Tagb. Moluccas: mandalirung'a, Talaud, 
pulton rupong, Banda, ritfut % Buru. West New 
Guinea: fulukwa, Manokwari, hornuu, Sorong; 

I ast New Guinea: komuktU, Mating name, ituttui. 



54 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 17. Young blukar (regrowth or secondary forest) on an abandoned tea estate near Tapos, West Java, 
c. 1000 m, consisting of three layers: 2 m high stand of Eupatorium inulifolium, 5-6 m high tree ferns of 
Cyathea contaminans, above which is an open canopy of Trema orientalis (L.) Bl. (Photogr. van Steenis). 



Kainantu, seraun, Daga-Bonenau, wanip, Enga 
lang., wantip, Medlpa, Wahgi, warium, Mendi. 

4. Trema angustifolia (Planch.) Bl. Mus. Bot. 2 
(1856) 58; Hook. /. Fl. Br. Ind. 5 (1888) 484; 
Gagnep. Fl. Gen. I.-C. 5 (1927) 686; Hand.-Mazz. 
Symb. Sin. 7 (1929) 108. — Sponia angustifolia 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 326; Miq. Fl. 
Ind. Bat. 1, 2 (1859) 215; Planch, in DC. Prod. 17 
(1873) 202. — Sponia acuminatissima Miq. Sumatra 
(1861) 410; Planch, in DC. Prod. 17 (1873) 202. — 
Sponia sampsonii Hance, Ann. Sc. Nat. V, 5 (1866) 
242. — T. acuminatissima (Miq.) Boerl. Handl. 3 
(1900) 358. — T. lanceolata Merr. Lingn. Sc. J. 7 
(1931) 302. — T. sampsonii (Hance) Merr. & 
Chun, Sunyatsenia 5 (1940) 40. — T. orientalis var. 
bicolor De Wit, Bull. Bot. Gard. Btzg III, 18 (1949) 
190. 

Shrub or small tree with spreading and drooping 
branches, 3-7 m, 5-15 cm 0. Branchlets densely set 
with rufous multicellular glandular hairs and 
glaucous short and matted 1 -celled hairs, sub- 
glabrescent. Stipules linear-lanceolate, 3-4 by 
1 mm. Leaves chartaceous to thin-coriaceous, 
narrow ovate-lanceolate to lanceolate, (3-)5-8(-10) 
by (l-)2-3(-4) cm, index 3-3V 2 , broadest below or 
at the middle ; discolorous, upper surface strongly 
scabrate, dark chocolate-brown to blackish-brown, 



lower surface densely set with short and matted 
rufous to glaucous 1 -celled and multicellular 
glandular hairs; base rounded to attenuate, sym- 
metrical; margin finely serrate throughout; apex 
acute to acuminate; midrib and nerves slightly 
raised beneath, impressed above; nerves 4-5 pairs, 
straight, ascending at 30-40°; reticulations fine, 
subscalariform to subareolate, obscure above and 
faintly visible beneath; petiole (2-)3-6(-8) by 
1 mm, densely short hairy. Inflorescences c? or c??, 
densely set with a short rufous indumentum, much- 
branched, (5-)10-15(-30)-flowered, at anthesis 
condensed, shorter than or as long as the petiole; 
bracts narrow ovate-acute, c. 1 l 2 -\ by V4-V2 mm. 
— S Flowers c. \-\ l l 2 mm 0; perianth lobes 5, 
elliptic, c. 1-1 */a by 1 mm; filaments c. 1 mm long, 
c. l / 2 -l mm ; pistillode obovoid-ellipsoid, strong- 
ly compressed, c. V2-I by V4 mm. — $ Flowers 
ovoid-conical, c. 2 by 1 mm; perianth lobes 5, 
narrow-lanceolate, acute, V2-I by 1 U- 1 l 2 mm; 
staminode absent; ovary c. 1-2 by 1 mm; stig- 
matic arms c. l / 2 -l mm, spreading or incurved. 
Drupe c. \ x \ 2 -1 mm 0, turning orange to red when 
ripe. Endosperm copious. 

Distr. China (Yunnan, Hainan), Thailand, and 
Indo-China; in Malesia: Malay Peninsula (com- 
mon), Sumatra (rare), Borneo (rare), Celebes (very 
rare). 



1977] Ulmaceae (Soepadmo) 55 

Ecol. Scattered in newly available habitats in Excluded 

the lowlands to submontane regions, from sea-level t tD ■ <. , , r ,. n , ixxin 

to 1200 m. FL fr. Jan.-Dec. Ripe fruits are dis- R ^T^T^Q?^ LV"^ / rr' 

persed by various species of bulbils. ^,£t ¥?JJ}?%} ? l = D * bre S easia longifoha 

Vern. Malaya: menarong, mengkirai, M; Wvrm. f.) Wedd. (Urt.caceae). 
Sumatra: kayu anggurung, M, Eastcoast, mang- 
kirai ketjil, M, Palembang. 

4. CELTIS 

Linne, Gen. PI. ed. 5 (1754) 467; Sp. PI. 2 (1753) 1043; Planch. Ann. Sc. Nat. Ill, 
10 (1848) 262; Bl. Mus. Bot. 2 (1856) 70; Miq. Fl. Ind. Bat. 1, 2 (1859) 220; 
Planch, in DC. Prod. 17 (1873) 168; B. & H. Gen. PI. 3 (1880) 354; Engl, in E. & 
P. Nat. Pfl. Fam. 3, 1 (1888) 63; Bernard, Bull. Herb. Boiss. II, 5 (1905) 1112, 
maps 9-15; J. J. Smith in K. & V. Bijdr. 12 (1910) 639; Leroy, Fl. Madag. et Com. 
Fam. 54 (1952) 3; Polhill, Kew Bull. 19 (1964) 139; Hutch. Gen. Fl. PI. 2 (1967) 
147; Elias, J. Arn. Arb. 51 (1970) 32; Soepadmo in Whitmore, Tree Fl. Mai. 2 
(1973) 414. — Solenostigma Endl. Prod. Fl. Norf. (1833) 41; Bl. Mus. Bot. 2 
(1856) 66; Miq. Fl. Ind. Bat. 1, 2 (1859) 219. — Fig. 18, 20, 22-23. 

Small to large monoecious or polygamo-monoecious trees, often buttressed. 
Bark smooth or finely fissured, often conspicuously warty lenticellate. Branches 
(in Mai.) unarmed, initially densely yellow-brown or rufous-hairy, glabrescent; 
hairs 1 -celled. Buds enclosed by the overlapping stipules or naked. Stipules thick 
and tough, peltately attached or free and scarious, caducous. Leaves entire dt not, 
3-nerved at base, semideciduous or persistent. Inflorescenes <$, ^, or 6**, branched 
racemes or panicles, few- to many-flowered, axillary or subterminal on the new 
shoot; staminate inflorescences borne on the lower and leafless part or in the axil of 
leaves of the new shoot; in the g't inflorescence the ^ flowers are borne on the 
distal ends of the axes; bracts minute, caducous. — Staminate (S) flowers globular, 
pedicelled or sessile; perianth lobes 4-5, imbricate in bud, membraneous, boat- 
shaped, outside sparsely pubescent, at anthesis recurved, caducous; stamens 
glabrous, inserted on the densely pilose receptacle; filaments subulate, incurved in 
bud and spreading elastically, exserted at anthesis; anthers ovoid to subreniform, 
dorsifixed just above the emarginate base, extrorse; pistillode present or absent. — 

I lowers ovoid, pedicelled; perianth lobes 4-5, imbricate in bud, connate at base, 
membranous, outside sparsely pubescent, boat-shaped, at anthesis recurved, cadu- 
cous; stamens well-developed and functional or rudimentary, other characters as in 
3 flowers; ovary ovoid-ellipsoid, sessile, style short or ± absent; stigmatic arms 
elongate, divergent, the tips entire to deeply bifid; ovule anatropous. Drupe fleshy, 
ovoid, ellipsoid or globose; exocarp thick and firm, mesocarp thin and fleshy, con- 
taining slimy substances; endocarp hard and persistent, smooth or variously ridged 
or pitted. Seed: coat membranous, chalazal area broad, dark-coloured and close to 
the minute hilum ; endosperm scanty or wanting, oily or gelatinous, nearly enclosed 
between the folds of the cotyledons. Embryo curved, cotyledons broad, foliaceous, 
equal or unequal in thickness, flat or conduplicatc, variously folded, incumbent on 
or embracing the short superior and ascending radicle. Germination epigeal. 

Dittr. About 30 W) spp ., widely distributed in tropical and temperate regions of the w.orld, the majority 
of species (30 40) in the Old and New World tropics, throughout \talc\iu ( { ) spp.). I ig. 19, 21. 



56 



Flora Malesiana 



[ser. I, vol. 8 2 




x 8 10 8 f %!Z Ph ! l WTi S B , LANCa a - Ne I w c shoot bearing flowers, x »/* b-c. $ flowers, x 8, d-e. 6 flowers 

L'Y', frUltS ' 1 /3,g : fruit, exocarp in LS, x 1V 3 , h. stone, endocarp in LS, x IV, i seed showing testa 

and chakza nat. size, j ditto in LS showing folded cotyledons, shaded oily endosperm, xl V 3 ?k embryo 

' fl J ?' ~ C \k emnd !? RoxB - U New shoot bearing flowers, x V m-n 5 flowers x 8 oT 

flowers, x 10 (a-c Merrill 52, </-e BS 1920,/-* bIrtlett 15071, /-Tschmu?z 16661 



1977] 



Ulmaceae (Soepadmo) 



57 




Fig. 19. Approximate range of Celtis L. Fossil records are indicated by dots. 



Fossils. Numerous fossilized wood-fragments, leaf-impressions, drupes and pollen grains have been 
discovered in various localities in Europe, the U.S.A., and in Asia. According to Elias I.e. the first 
(oldest) records are apparently from the early Eocene in Wyoming and the late Eocene in Georgia, U.S.A. 
Continuing through the Oligocene, Celtis spp. are best known from the Miocene, fossilized material of 
younger age is relatively sparse at least in the U.S.A. — References: A. Graham (ed.), Floristics and 
Paleofloristics of Asia & Eastern North America (1972) 147; Greguss, Tert. Angios. Hung., Ak. Kiado 
Budapest (1969) 83; La Motte, Mem. Geol. Soc. Am. 51 (1952) 112. Fig. 19. 

Ecol. Malesian species of Celtis may be classified into two rather distinct ecological groups, i.e. the 
C. philippensis var. philippensis and C. tetrandra groups. The first group, which includes also C. hilde- 
brandii, C. latifolia, C. luzonica, C. paniculata, and C. rigescens, is found mainly in the lowland forests, 
both primary and secondary, and is an important constituent of the understorey tree community in moist 
areas. The C. tetrandra group, which includes C. rubrovenia, C. timorensis, and C. philippensis var. wightii, 
is confined to areas which are subject to a rather pronounced seasonal climate, or if they occur in wetter 
regions, they grow on strongly drained substrates, e.g. rocky shores, limestone, etc. In tune uith this 
environmental preference, the second group shows a more prominent flush-wise growth habit and its 
species are completely or partly deciduous. In Malaya, species of Celtis are producing flowers around 
July-September, while fruit ripens January-March. 

How pollination is affected is not certain, but judging from the structure and position of the inflores- 
cence, some species (e.g. C. philippensis var. philippensis, C. hildebrandii, and C. latifolia) may be pollinated 
by insects, while others (C. tetrandra, C. timorensis, C. rubrovenia, and C. rigescens) may be pollinated 
by wind. 

The ripe fleshy drupes which turn to orange, red or bluish-black may be dispersed by birds, or alter- 
natively they may be dispersed by water as the embryo is protected by the hard, persistent and durable 
endocarp. 

Morph. Except for C. paniculata the stamens of the pistillate flowers are well-developed and functional. 
In the male or staminate flowers the pistillode is rudimentary or completely absent in C. tetrandra, 
C. timorensis, and C. rubrovenia. In the other species the pistillode is present and relatively rather well- 
developed though non-functional. 

Chromosomes. The chromosome numbers reported are: n = 10 (2n = 20) (C. australis var. eriocarpa, 
C. inguana, C. laevigata, C. occidentalis, ( . sinensis, and C. timorensis (under C. cinnamomea)); 2n = 22 
(C. spinosa );2n = 28(C.occidentalis);2n = 40 (C. australis and C. tupalangi). — References: Sax, J. Arn. 
Arb. 14 (1933) K2; Bowdin, Am. J. Bot. 32 (1945) 195; Dari.in<, ion & WYLIE, (hromos. Atlas (1955) 
182; Mehra & GILL, Taxon 17 (1968) 574; Gadella et al. Acta Bot. Neerl. 18 (1969) 74; Mehra & Hans. 
Taxon 18 (1969) 310; FEDOROV (ed). (hromos. Numbers Flow. PI. (1969) 710; Mehra & Gill, J. Am. 
Arb. 55 (1974) 663. 

According to Sax I.e. there seems to be at least in C. occidentalis a high degree of pollen sterility and a 
high incidence of meiotic irregularity. This may be one of the causes why in Celtis there is a very high 
percentage of barren seeds production, even among tropical species. 

■ I detailed study on the microsporogencsis, CTCgaspO f Og BnCMI and embryogencsis of 
Ctltix species has ever been carried out. In Malcsia the solitary ovule is bitegmic, anatropous and inserted 
just below the apex of the loculc. After fertilization both integuments develop into thin membranous 
seed coats with a broad, dark-coloured, more or less circular chala/a. I he endocarp becomes woody and 
very hard and impregnated by mineral deposits. It is persistent and becomes variously sculptured (ridged, 



58 Flora Malesiana [ser. I, vol. 8 2 

pitted, or nearly smooth). The embryo is strongly curved with the hypocotyle superior and ascending, 
situated in between or nearly enclosed by the broad, thick, foliaceous cotyledons. The cotyledonar lobes 
are somewhat unequal in thickness, and they are either induplicate or variously folded. Endosperm is very 
scanty to absent and either gelatinous or oily. Especially in C. paniculata and C. tetrandra, at least 70-80% 
of the fruits produced are barren. Though the fruits are developed normally, the embryo fails to grow and 
becomes shrivelled. As a result the fruits are empty. 

KEY TO THE SPECIES 

(Measurements of leaf and fruit based on fully mature material) 

1. Leaves entire or nearly so. <$ Inflorescence a much-branched many-flowered panicle with up to 150 
flowers. Pistillode rather well-developed, c. I-lVa by V2-I mm. Stigmatic arms bilobed or bifid at the 
tip. 
2. Leaves rugose, brittle, sparsely pubescent beneath; midrib and lateral nerves strongly raised beneath. 
Stipules not peltately attached, free from one another. Fruit densely appressed-hairy. 1. C. rigescens 
2. Leaves not rugose, not brittle, glabrous; midrib and nerves only slightly raised beneath. Stipules 
peltately attached, overlapping. Fruit glabrous. 
3. Leaves with (2-)3-5 pairs of nerves. 

4. Leaves elliptic-orbicular or elliptic-oblong, index (lV4-)lV2(-2); midrib and nerves slightly raised 
beneath; lowest pair of nerves running to 2 / 3 - 3 / 4 the length of the leaf, upper pairs of nerves ascend- 
ing and arcuating. Stigmatic arms shallowly bilobed at the tip. Fruit globose, 16-20 by 14-18 mm. 

2. C. luzonica 
4. Leaves elliptic or ovate-eliptic, index (l 1 j 2 -)2-2 1 l 2 (-'i)\ midrib and nerves flattish beneath; lowest 
pairs of nerves running up to Va-VaC-^/s) the length of the leaf; upper pairs of nerves weak, sub- 
horizontal. Stigmatic arms deeply bifid at the tip. Fruit ovoid or ellipsoid, 7-12 by 5-8 mm. 
5. Nerves 3-5 pairs; lowest pair running to V3-V2 the length of the leaf. Inflorescence mostly 3 or $. 
Stamens of $ flower rudimentary, non-functional. Fruit ovoid, feebly 4-5-angular in CS; endo- 

carp with reticulate ridges 3. C. paniculata 

5. Nerves 1— 2(— 3) pairs; lowest pair running up to 2 / 3 the length of the leaf. Inflorescence 6* or cJ5- 
Stamens of $ flower well-developed and functional. Fruit ellipsoid, ± terete; endocarp smooth. 

4b. C. philippensis var. wightii 
3. Leaves with 1 pair of nerves. 
6. Leaf symmetrical. Inflorescence <$ or c?$. 6* Inflorescence 15-40-flowered. Cotyledons curved but 

not folded. 
7. Leaves (4— )8-14(-18) by (2-)3-6(-8) cm; reticulations fine, dense. Ovary glabrous. Fruit 8-15 by 

7-12 mm; endocarp smooth 4a. C. philippensis var. philippensis 

7. Leaves (8—) 1 5—1 8(— 25) by (6— )8-12(— 18) cm; reticulations coarse, wide-spaced. Ovary densely 

appressed-pubescent. Fruit 15-25 by 10-18 mm; endocarp pitted 5. C. latifolia 

6. Leaf oblique. Inflorescence <J and g. d> Inflorescence 60-150-flowered. Cotyledons curved and many 

times folded 6. C. hildebrandii 

1. Leaves serrulate to denticulate at least in the upper half. S Inflorescence racemose, 5-20-flowered. 

Pistillode strongly reduced in size or absent. Stigmatic arms entire. 
8. Leaf not strongly oblique in outline; nerves 1— 2(— 3) pairs. Inflorescence <3 and <$$; <$ flower 5-merous. 
9. Leaves (6-)10-12(-17) by (2 1 1 2 -)4-5(-8) cm, index \ l l 2 -2 l l 2 . 6* Inflorescence 10-20-flowered. Ovary 
glabrous. Infructescence 4-5 cm long. Fruit ovoid, strongly beaked, 5-10 by 3-6 mm. 

7. C. timorensis 
9. Leaves (3-)4-5(-67 2 ) by (l 1 /4-)l 1 /2-2 1 /2(-3 1 / 2 ) cm, index 2-3.3. <$ Inflorescence 5-7-flowered. Ovary 

densely hairy. Infructescence c. IV2 cm. long. Fruit globose, not beaked, c. 3 by 3 mm 

8. C. rubrovenia 
8. Leaf strongly oblique ; nerves 3-4 pairs. Inflorescence <$ and $ ; $ flower 4-merous. . 9. C. tetrandra 

1. Celtis rigescens (Miq.) Planch, in DC. Prod. 17 and sparsely minute lenticellate. Terminal buds 
(1873) 182; Soepadmo in Whitmore, Tree Fl. Mai. ovoid-conical, c. 3 by 2 mm, scales densely yellow- 
2 (1973) 416. — Solenostigma rigescens Miq. ish-brown tomentose. Stipules linear-lanceolate, 
Sumatra (1861) 411. — Solenostigma sumatrana 4-5 by 1-1 V2 mrn- Mature leaves thick-coriacecus, 
Miq. I.e. 411. — C. sumatrana (Miq.) Planch, in strongly rugose, stiff and brittle when dry, ovate- 
DC. Prod. 17 (1873) 181. — C. nymanii K.Sch. in elliptic to elliptic-oblong, (5-)8-12(-15) by (2V2-) 
K.Sch. & Laut. Fl. Schutzgeb. Nachtr. (1905) 3-5(-6V 2 ) cm (index l'/a^Va). broadest at or 
240; Laut. Bot. Jahrb. 50 (1913) 311. — C. asperi- slightly below the middle; above glabrous, shining, 
folia Merr. Philip. J. Sc. 17 (1920) 246; En. Philip, beneath sparsely yellowish-brown pubescent es- 
2 (1923) 32. — Fig. 20e, 23a-b. pecially on midrib and nerves; base rounded to 
Large tree up to 45 m, 1 m 0. Buttresses up to subcordate, symmetrical, rarely attenuate-rounded 
6m tall, 3 m out, 5cm thick. Bark grey-brown, and slightly asymmetrical; margin undulate, en- 
smooth, finely fissured to pustulate and lenticellate. tire or distantly serrate in the upper half, very often 
Innovations densely yellowish brown to rufous incurved; apex rounded-acute to acuminate; 
simple hairy. Older twigs glabrous, finely striate midrib and nerves strongly raised beneath, flattish 



1977] 



Ulmaceae (Soepadmo) 



59 




Fig. 20. Celtis paniculata (Endl.)Sp\ch. a. 9 Inflorescence, x 8, b. flower, x 14, c. fruit, x l'/s.d. shoot 
apex showing terminal bud above 2 pairs of overlapping stipules, x 5. — C. rigescens (Miq.) Planch. 
e. Embryo, folded cotyledons, x Vj 3 . — C. timorensis Span. f. Embryo in LS, x 2. — C. hildebrandii 
Soepadmo. g. Pitted endocarp, x 3, h-i. embryos, x 4. — C. latifolia (Bl.) Planch, j. Fruit, nat. size 
(a-b Kornassi 463, c. Kostermans & Soegeng 352, d Pleyte 73, e Bloembergen 4231, /Forbes 1073, 

g-i Clemens 8375, j BSIP 1 1768). 



to deeply impressed above; main nerves (2-)3(-4) 
pairs, ascending and arcuating, anastomosing 
along the margin, the lowest pair running up to 
1 2- 3 4 the length of the leaf; reticulations sub- 
scalariform to subareolate, distinct beneath; 
petiole 6-10 by 1-2 mm, shallow-sulcate, densely 
appressed yellowish-brown pubescent, glabrescent. 
Inflorescence $ or * or rarely mixed, 1 l l 2 -5 cm long; 
bracts ovate, c. l ! 2 -l by Vi mm. — 3 Inflorescence 
a much-branched, multiflorous (up to c. 70) 
panicle up to 5 cm long, axes slender, flexuous, 
borne on the lower and leafless part of the new 
shoot or in the axils of lower new leaves; <$ flower 
1 ' j-2 mm 0, subsessile ; perianth lobes (4-)5, elliptic, 
c. 1-1 ' 2 by 1 mm; filaments I-l'/^mm long, anthers 
reniform, c. 1 by '/imm; pistillode minute, sur- 
rounded by dense pale yellowish-brown hirsute 
hairs. In a mixed inflorescence, the few § flowers 
are situated near the tip of the axis. — $ Inflores- 
cences borne in the axils of leaves of the new shoot, 
1 I' 2 cm long with up to 5 flowers; 5 flower 
ovoid, c . 2 by 1 mm; perianth lobes (4-)5, elliptic- 
rounded, c. f'/j by 1 mm, stamens (4 )5; filaments 
c. 1 mm, anthers r. '/i by J /i mm; ovary slightly 
compressed, densely pale yellowish-brown ap- 
prcsscd-hairy ; stigmatic arms spreading, bifurcate 
at the tip. Infructescence up to 2 3 cm long, axes 
up to 2mm thick, with (I )2( 3) fruits. Fruit 



ellipsoid, faintly 5-angular, c. 15 by 10 mm, 
appressed-pubescent, glabrescent ; exocarp 2-3 mm 
0, rather woody, occasionally lenticellate, turning 
deep-red when ripe, containing slimy substances 
when boiled. Embryo curved, cotyledons folia- 
ceous, folded, equal. Endosperm scanty. 

Distr. Solomons (common); in Malesia: New 
Guinea (incl. New Britain, common), Moluccas 
(Ceram, Buru, Sula, Morotai), NE. Celebes (Mina- 
hasa), E. Borneo (W. Kutei), W. Java (Bantam), 
Central & S. Sumatra, Anambas Is., and Malaya 
(Perak, Selangor, Pahang, Johore). 




Fig. 21. Species density of Celtis L. in Malesia; 
above the hyphen endemic spp., below it the non- 
endemic ones. 



60 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 22. Celtis philippensis Blanco var. wightii (Planch.) Soepadmo at Atasangin, Bodjonegoro (Photogr. 

Wind, Jan. 1925). 



1977] 



Ulmaceae (Soepadmo) 



61 



Ecol. In primary and secondary forest, 
0-1500 m, on various types of soils including lime- 
stone. Often rather common locally (W. New 
Guinea and Solomons). Fl. (March-April) Sept., 
Dec.,/r. Dec.-Aug. 

Vern. Malay Peninsula: mimpilas bulan, Abor., 
rimpilas, M. Sumatra: klutum, marsikan, rim- 
pilas, M. Palembang, asin asin, Anambas. West 
New Guinea: sihiega, siriega, Manikiong lang., 
wennimaram, Berik lang., Hollandia, bigek, 
Oransbari, Hattam lang., piehQi), Kebar, bipa, 
Wandammen lang., Adi I., ainam, Key lang., 
hajanggwij, manji, sikika, Manokwari, pimar, 
Sidei lang., warin, siempai, wiempai, Biak lang., 
walik, walis, Mooi lang.; East New Guinea: 
gairama, watot, Waria dial., Moresby, Madang 
Distr., koru, Bambi, suri, Madang, bagibib, 
Kaigorin, aban, bison, sam, Bilia, baigu(p), Amele, 
bagibip, kara, sungung, susuik, Dumpu, goga, 
mutum, sawan, Faita, tapang, Washuk, Sepik 
Distr. ; ailee, Urin, S. New Britain ; gwalafalisi, 
lausi, Kwara lang., Solomons. 

Note. Sterile specimens, especially those with 
young leaves only, may easily be mistaken as 
belonging to Grewia or Microcos (Tiliaceae) or to 
Ziziphus angustifolius (Bl.) Hatus. 

2. Celtis luzonica Warb. in Perkins, Fragm. Fl. 
Philip. (1905) 164; Merr. En. Philip. 2 (1923) 32. 
— Fig. 23c. 

Tree up to 30 m, 90 cm 0. Bark smooth yellow- 
ish-grey. Innovations sparsely set with minute 
simple hairs, glabrescent. Terminal bud ovoid, c. 6- 
10 by 3-5 mm. Stipules ovate, thick, glabrous, 
peltately attached and overlapping, c. 5-10 by 3-5 
mm. Leaves broad elliptic to elliptic-oblong, thick- 
coriaceous, glabrous, (8-)12-16(-19) by (5-)8-10 
(-12) cm, index (l 1 / 4 -)l 1 / 2 (-2); base rounded, 
symmetrical; margin entire, undulate, apex 
rounded with blunt and sometimes emarginate tip; 
midrib and nerves raised beneath, flattish to shal- 
lowly impressed above; nerves 2-3 pairs, ascending 
and arcuating, at least at an angle of 55 = with the 
midrib, anastomosing and looped along the margin, 
the lowest pair running up to 2 3 - 3 4 the length of 
the leaf; reticulations irregular to subscalariform, 
sparse, distinct beneath or obscure on both sur- 
faces; petiole 10-20 by lV 2 -2(-3) mm, flat or 
shallowly sulcate. Flowers 5-merous. Inflorescences 
o or 4 'i, much-branched panicles or racemose, up 
to 150-flowered, up to 10 cm long, axillary or sub- 
terminal, including the bracts sparsely puberulous, 
glabrescent; bracts ovate, minute, c. 1 by l l 2 mm. 
lower c. 1 ' 2 -2 mm ; perianth lobes c. 1 1 / 2 - 
2 by 1 mm; filaments l 1 2 -2 mm, anthers subreni- 
form, c. 1 — 1 * / 2 by \ 2 -l mm; pistillode ovoid- 
conical, compressed, c. J by ', 2 mm. — 5 Flowers 
ovoid-conical, borne at the distal parts of the 
inflorescence, 2'/ 2 -3 by 2-2'/ 2 mm; perianth lobes 
r. 1' 2 2 b* 1 l' 2 mm; filaments c. l-l' 2 mm, 
anthers c. ! 4 by '/ 2 mm; ovary | compressed, 
sparsely pubescent apically, c. 2-2'/ 2 by 1-1 '/j mm; 
stigmatic arms spreading or incurved, broadened 
and shallowly bifid at the tip. Infructescence with a 
stout axis 2-3 mm thick, bearing up to 10 fruits. 
Fruit globose, glabrous, 16-20 by 14-18 mm, | 
terete, reddish-brown when ripe; endocarp 4-lobed 
and splitting at maturity; embryo curved, hypoco- 
tyle ascending, cotyledons foliaccous, equal, 



folded ; endosperm very scanty to nearly absent. 

Distr. Malesia: Philippines (Luzon, Surigao, 
Mindanao, Mindoro, Samar, Masbate). 

Ecol. In thickets and forests at low altitude. 
Fl.fr. March- July. 

Vern. Philippines: malaikmo, malaitmo, Tag., 
daloo, Todaya. 

3. Celtis paniculata (Endl.) Planch. Ann. Sc. 
Nat. Ill, 10 (1848) 305; in DC. Prod. 17 (1873) 182; 
Bth. Fl. Austr. 6 (1873) 156; Nadeaud, En. PI. 
Tahiti (1873) 42; Laut. Bot. Jahrb. 50 (1913) 310; 
Kaneh. Fl. Micron. (1933) 82, f. 14; Francis, 
Austr. Rain-forest Trees ed. 2 (1951) 67. — Sole- 
nostigma paniculatum Endl. Prod. Fl. Norf. 
(1833) 42. — C. pacifica Planch. Ann. Sc. Nat. Ill, 
10 (1848) 308; in DC. Prod. 17 (1873) 184. — 
C. ingens F.v.M. Fragm. 4 (1864) 88. — C. vitiensis 
A. C. Smith, Bull. Torr. Bot. CI. 70 (1943) 536; 
J. Arn. Arb. 31 (1950) 150; Parham, PI. Fiji Isl. 
(1964) 88. — Fig. 20a-d, 23i. 

Small to large tree, up to 36 m, 70 cm 0. 
Buttresses occasionally present, up to l l J 4 m. Bark 
smooth to finely fissured, light- to dark-brown. 
Innovations sparsely appressed-puberulous, glab- 
rescent. Older twigs glabrous, sparsely lenticellate. 
Stipules overlapping, embracing the twig and 
enclosing the bud, broad-ovate-acute, thick, 
c. 4-5 by 3-4 mm. Leaves elliptic to ovate-elliptic, 
(5-)8-ll(-13) by (2V 2 -)3-5(-6) cm, index (lV 2 -)2 
(-2V2) ; coriaceous, glabrous, dull grey-green when 
dried ; base attenuate-rounded, mostly symmetrical, 
margin entire, often undulate, apex bluntly acute 
to rounded-acute; midrib and nerves only slightly 
raised beneath, flattish to shallowly impressed 
above; main nerves 3-5 pairs, ascending and ar- 
cuating at an angle of over 50° with the midrib, 
anastomosing and looped along the margin, the 
lowest pair running up to V3-V2 the length of the 
leaf; reticulations fine, irregular, obscure on both 
surfaces; petiole 6-15 by 1 —1 1 / 2 mm, flat to ± sul- 
cate. Flowers 5-merous. Inflorescences 6* or $ or 
rarely mixed, axillary or subterminal, much- 
branched, 5-60-flowered, including the bracts 
sparsely appressed-puberulous; bracts ovate-acute, 
c. 2 by 1 mm. — <S Inflorescences up to 4 cm long, 
borne on the lower and leafless part or axillary on 
the lower leaf of the new shoot, paniculate, 30-60- 
flowered; o flowers subsessile, c. l-l'^mm 0; 
perianth lobes transparent, c. 1-lVa by 72 mm; 
filaments c. 1 mm, anthers subreniform, c. 3 / 4 -l by 
1 2 mm; pistillode minute, compressed. — Mixed 
or 4 inflorescences axillary on the upper parts of the 
new shoot, racemose, 5-10-flowered; 9 flower 
ovoid-conical, c. 2 by 1 mm; perianth lobes ovate- 
acute, c. V2-I by \ 2 mm, subglabrous; staminodes 
rudimentary, very much shorter than the perianth; 
ovary ± compressed, subglabrous, c. l'/ 2 -2 by 
1 mm; stigmatic arms spreading, bifid at the tip. 
Fruit ovoid, faintly 4-5-anguIar, beaked, glabrous, 
7-12 by 5-8 mm, bluish to glaucous when ripe, 
mostly sterile. 

Distr. Australia (Queensland, Norfolk I.), 
Melanesia (Solomons, New Hebrides, New Cale- 
donia), Polynesia (Fiji, Tonga, Tahiti, Cook Is., 
Pitcairn, Mangarawa I., Tuamotus, Marquesas), 
Micronesia (Marianas); in Malesia'. I esser Sunda 
Islands (Wetar), Borneo (Mt Kinabalu, very rare), 
Moluccas (Morotai, Ceram, Ambon, Tanimbar), 



62 



Flora Malesiana 



[ser. I, vol. 8 2 



New Guinea (common in West, apparently rare in 
East). 

Ecol. Primary and secondary forest, 0-900 m, 
on well drained soils including coral limestone, 
very often common locally (W. New Guinea and 
Solomons). Fl. fr. July-May. 

Vern. New Guinea: wiempai, Biak, sehiega, 
Manikiong; Solomons: lausiasi, Kwara; Fiji: 
marasa, Sabalu. 

4. Celtis pbilippensis Blanco, Fl. Filip. (1837) 197; 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 306; in DC. 
Prod. 17 (1873) 184 ( i philippinensis , )\ Bth. Fl. 
Austr. 6 (1873) 156; Vidal, Rev. Pl. Vase. Filip. 
(1886) 248; Merr. Philip. J. Sc. 1 (1906) Suppl. 42; 
Sp. Blanc. (1918) 122; En. Philip. 2 (1923) 32; 
Leroy, Bull. I.F.A.N. 10 (1948) 212, incl. var. 
consimile (Bl.) Leroy; Fl. Madag. Fam. 54 (1952) 
3; Soepadmo in Whitmore, Tree Fl. Mai. 2 (1973) 
416. — C. wightii Planch. Ann. Sc. Nat. Ill, 10 
(1848) 307; Wight, Ic. PI. (1853) t. 1969; Planch. 
in DC. Prod. 17 (1873) 184; Hook./. Fl. Br. Ind. 5 
(1888) 483; Brandis, Ind. Trees (1906) 594; J. J. 
Smith in K. & V. Bijdr. 12 (1910) 647; Gagnep. 
Fl. Gen. I.-C. 5 (1927) 683 ; Polhill, Kew Bull. 19 
(1964) 141 ; Back. & Bakh./. Fl. Java 2 (1965) 11. 
— C. strychnoides Planch. Ann. Sc. Nat. Ill, 10 
(1848) 306; in DC. Prod. 17 (1873) 185; Warb. 
Bot. Jahrb. 13 (1891) 287; Laut. in K.Sch. & Laut. 
Fl. Schutzgeb. (1900) 264. — C. mauritiana 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 307; in DC. 
Prod. 17 (1873) 184. — Sponia strychnifolia 
Teysm. & Binn. Nat. Tijd. N. I. 4 (1853) 394; Ned. 
Kruidk. Arch. 3 (1855) 392. — Solenostigma 
brevinerve Bl. Mus. Bot. 2 (1856) 67. — Solenos- 
tigma laurifolium Bl. I.e. 68 ; Miq. Fl. Ind. Bat. 1 , 2 
(1859) 220, incl. var. constricta Miq. — Solenos- 
tigma hasseltii Bl. Mus. Bot. 2 (1856) 68. — Sole- 
nostigma consimile Bl. I.e. 68. — Solenostigma 
djungiel Bl. I.e. 69. — Solenostigma philippinensis 
(Blanco) Miq. Fl. Ind. Bat. 1, 2 (1859) 220. — 
Solenostigma wightii (Planch.) Miq. I.e. 220. — C. 
brevinervis (Bl). Planch, in DC. Prod. 17 (1873) 
183. — C. laurifolia (Bl.) Planch. I.e. 185. — 
C. hasseltii (Bl.) Planch. I.e. 185. — C. djungiel 
(Bl.) Planch. I.e. 185. — C. mindanaensis Elmer, 
Leafl. Philip. Bot. 8 (1915) 2842. — C. collinsae 
Craib, Kew Bull. (1918) 370; Ridl. Fl. Mai. Pen. 3 
(1924) 322. — C. multifolia Elmer, Leafl. Philip. 
Bot. 10 (1939) 3796, angl., inval. — Fig. 18a-k, 22. 
Small to large tree, up to 30 m, 80 cm 0. 
Buttresses if present up to 2V2 m tall, 2 m wide and 
10 cm thick. Bark smooth to finely fissured, pale 
grey to grey-brown. Innovations initially sparsely 
to densely set with yellowish-brown appressed 
or/and woolly hairs, glabrescent. Stipules ovate- 
acute, 6-10 by 2-4 mm, thick, peltately attached, 
overlapping and enclosing the bud. Leaves thick- 
coriaceous, glabrous, full grey when dried, elliptic- 
oblong to suborbicular, (4-)8-14(-18) by (2-) 
3-6(-8)cm, index (lV 2 -)2-3; base rounded or 
attenuate-rounded, mostly symmetrical; margin 
entire often undulate (immature leaves very rarely 
distantly serrulate at the upper half) ; apex rounded 
to acute; midrib and nerves raised beneath, im- 
pressed to flattish above; main nerves 1 pair, 
ascending, arcuating and running throughout the 
length of the leaf (var. philippensis) or 1-3 pairs, 
the lowest pair ascending, arcuating, and running 



up to about 2 / 3 the length of the leaf and then 
anastomosing with the 1-2 weaker and more or 
less horizontal upper nerves (var. wightii) ; reticula- 
tions fine, dense, subscalariform or subareolate, 
usually rather distinct beneath; petiole 6-15 by 
1-2 mm, sulcate. Inflorescences <$ or c?$, much- 
branched panicles, many-flowered, including the 
bracts densely yellow-brown to rufous soft-hairy; 
bracts ovate-acute, c. 3 by 1 mm. In the $ $ in- 
florescence the 5 flowers are borne on the distal 
parts of the inflorescence. — c? Inflorescences borne 
on the lower part of the new shoots, 2-4 cm long, 
with up to 40 flowers ; c? flowers c. 2 mm ; 
perianth lobes ovate-elliptic, c. V-\ 2 -2 by 1 mm; 
filaments l-l'^mm long, anthers subreniform, 
c. V2-I mm by x \ 2 mm ; pistillode ovoid-cylindrical, 
compressed, c. 1-1 1 / 2 by l j 2 mm. — Mixed in- 
florescence up to 5 cm long, up to 50-flowered, 
borne on the upper part of the new shoots; $ 
flowers ovoid, c. 2-2V 2 by 2 mm; perianth lobes 
ovate-elliptic, c. 2-2 1 l 2 by 1 mm; filaments 1-2 mm, 
anthers l j 2 -\ mm ; ovary ovoid-cylindrical, 
c. 2-2 1 l 2 by IV2-2 mm, glabrous except at the base; 
stigmatic arms spreading, c. 1-1 1 I 2 mm long, 
bilobed to bifid at the tip. Infructescence up to 
4-5 cm long, carrying 1-3 fruits, axes 1-2 mm 
thick. Fruit ovoid, glabrous, 8-15 by 7-12 mm, 
beaked when young; exocarp less than 1 mm 0, 
sometimes lenticellate, turning orange to red when 
ripe; endocarp ± smooth; embryo curved, hypo- 
cotyle ascending, cotyledons broad, foliaceous, 
unequal in thickness, not folded; endosperm oily, 
scanty to absent. 

Distr. Tropical Africa to Madagascar, Indian 
Ocean (Reunion, Mauritius, etc.), India, Burma, 
? SE. China, Hongkong, Taiwan, Indo-China, 
Thailand, throughout Malesia to NE. Australia 
and the Solomons. 

Taxon. A rather variable, widely spread species 
with two rather but not completely distinct 
varieties. These are: 

a. var. philippensis, characterized by larger 
leaves of (7— )9— 12(— 18) by 4-8 cm with one pair of 
nerves usually running more or less throughout the 
length of the leaf, subscalariform reticulation, and 
larger fruit of 10-15 by 8-12 mm; 

b. var. wightii (Planch.) Soepadmo, comb. nov. 
(basionym: C. wightii Planch. I.e. supra). Fig. 22. 
Characterized by smaller leaves, (4-)5-7(-9) by 
(2-)3-4(-5V2) cm with 1-3 pairs of nerves and the 
lowest pair mostly running up to 2 / 3 the length of 
the leaf, and slightly smaller fruit, 8-12 by 6-10 mm. 

It should be noted, that the distinguishing charac- 
ters mentioned above should be applied in com- 
bination; if taken individually they may not be 
clearly well defined. For example, there are several 
specimens (e.g. Gardner s.n., Thwaites CP 50 
from Ceylon; King s.n. and Browne s.n. from 
India; Parkinson 214 from the Andamans; 
Unesco 214 from Malaya; Jacobs 4709, 4711, and 
Kostermans 23061 from Java; Kostermans & 
Wirawan 61 from the Lesser Sunda Is.; NGF 
19100 & 30787 from New Guinea; Merrill Sp. 
Blanc. 52 from the Philippines, etc.) which have 
both types of venation. As for the fruit, the smaller 
size in var. wightii may be due in part to the fact 
that they are not fully ripe, as the majority of them 
are empty (without embryo). Furthermore, it was 
also noticed that most specimens of var. wightii 



1977] 



Ulmaceae (Soepadmo) 



63 



have been collected from localities under a strong 
seasonal climate or from trees growing on well- 
drained and poor soils (rocky or sandy beach, 
limestone hills, etc.). 

Ecol. Understorey tree in primary and secon- 
dary forests, on various types of soils, at low alti- 
tudes (0-650 m) ; often gregarious and very 
common locally. Fl. fr. mostly July-April,. The 
fruits which turn to orange or red when ripe may be 
dispersed by birds, but in the case of var. wightii, 
which mostly grows in the very coastal forest, they 
may be dispersed by sea-water as well ; (the endo- 
carp is woody, hard, and persistent). 

Uses. Though not durable, the wood is locally 
used for house-building. 

Vern. Java: ki-indog, ki-howe, S, W. Java, 
kiraja, pusutan, sintok, sipat, sepreh, tje~ngkek, 
wuluh, J, Central & E. Java; N. Borneo: nyelepi; 
Philippines : malaitmo, narabagsay, Tag. ; Celebes : 
kao lulu, Malili; Moluccas: horo, Morotai; 
Lesser Sunda Is. : menulang, Sumba, nemu, Flores; 
New Guinea: piih, Kebar, marmar, Tor, Berik, 
sehiega, Manikiong, bipiejit, Hattam, ware~n, 
Biak, ikai, ikoi, Kemtuk, milawar, Mooi, etc. 

5. Celtis latifolia (Bl.) Planch, in DC. Prod. 17 
(1873) 186; Warb. Bot. Jahrb. 13 (1891) 287; Laut. 
in K. Sch. & Laut. Fl. Schutzgeb. (1900) 264; Bot. 
Jahrb. 50 (1913) 311. — Solenostigma latifolium 
Bl. Mus. Bot. 2 (1856) 67; Miq. Fl. Ind. Bat. 1, 
2 (1859) 219. — Solenostigma zippelii Bl. Mus. Bot. 
2 (1856) 67. — C. zippelii (Bl.) Planch, in DC. 
Prod. 17 (1873) 186. — C. kajewskii Merr. & 
Perry, J. Arn. Arb. 22 (1941)254. —Fig. 20j, 23k. 
Tree up to 35 m, 80 cm 0. Buttresses plank-like, 
up to 2 m tall, 2\ 2 m out and 6 cm 0. Bark smooth 
to finely fissured, pustulate-lenticellate, light- 
brown to grey-brown. Innovations densely yellow- 
ish-brown hairy, glabrescent. Young twigs blackish 
when dry, older ones greyish, glabrous and sparsely 
lenticellate. Terminal buds ovoid-conical, acute, 
10-15 by 4-5 mm. Stipules peltately attached, over- 
lapping, thick, ovate-acute, c. 10 by 5 mm. Leaves 
thick-coriaceous, glabrous or sparsely pubescent 
beneath, especially on midrib and nerves, broadly 
ovate to elliptic-orbicular, (8-)15-18(-25) by 
(6-)8-12(-18) cm, index Vl 2 -2 l l 2 ; base rounded to 
subcordate, symmetrical to ± asymmetrical; 
margin entire, undulate, often recurved; apex 
bluntly rounded or acute to acuminate; midrib and 
nerves strongly raised beneath, impressed above; 
nerves 1-2 pairs, ascending and arcuating, the 
lowest pair running through •/« of the length of the 
leaf; reticulations coarse, wide-spaced, subscalari- 
form, distinct beneath; petiole 10-20 by 2-3 mm, 
glabrous, shallowly sulcate. Flowers 5-merous. 
Inflorescence J or o It, axillary or borne on the 
lower part of the new shoot, 10- 30- flowered, in- 
cluding the bracts densely yellowish-brown 
apprcssed-hairy; bracts ovate, c. 2 by 1 mm. — <$ 
Inflorescence (not fully developed) up to 2 cm long, 
15-30-flowered, paniculate; J flowers c. I 1 2 2 mm 
0; perianth lobes e. I ' 2 2 by 1 mm; filaments 
c. I mm, anthers <. ' 2 \ by '/ 2 mm; pUtillodc 
compressed ovoid, r. 1 by ' 2 mm. - J V Inflores- 
cence racemose, 5-10-flowercd, slender, up to 
7 cm long, few-branched; ',' flower ovoid-ellipsoid, 
borne on the distal part of the inflorescence, 
r. 2 3 by 2 mm; perianth lobes ovate-lanceolate, 



c. 2-2V2 by 1 mm, at anthesis recurved; filaments 
up to l l 1 2 mm, anthers c. 1 l 2 - 3 U by l l 2 mm; ovary 
ovoid-cylindrical, c. 2-3 by 1-1 72 mm, initially 
densely appressed-hairy, glabrescent except for 
the basal part ; stigmatic arms spreading, shallowly 
bilobed at the tip. Infructescence up to 5 cm long, 
carrying 1-5 fruits, axes sturdy c. 2-3 mm thick. 
Fruit ovoid, faintly 4-5-angular, glabrous, l 1 /*- 
2V 2 by l-l 3 / 4 cm, exocarp occasionally lenticellate, 
up to 2 mm 0, turning to orange or deep-red when 
mature. Embryo curved, cotyledons fleshy, un- 
equal in thickness, hypocotyle ascending; endo- 
sperm absent. 

Distr. Solomons (very common); in Malesia: 
Philippines (Palawan), Moluccas (Morotai, 
Tidore), and New Guinea (in West very common in 
the vicinity of Manokwari and Hollandia; in East 
in Sepik and Morobe Districts; New Britain). 

Ecol. Primary and secondary forests on sandy 
clay soils, 0-400 m. Fl. fr. mostly Jan.-Aug. 

Vern. Moluccas: tohu, Morotai; New Guinea: 
sehiega, Manikiong, marmar, Berik, bepiet, 
Hattam; Solomons: lae-lae, Kwara. 

6. Celtis hildebrandii Soepadmo, sp. nov. — Fig. 
20g-i, 231. 

Species valde affinis C. philippensi var. philippensi 
et C. latifoliae, sed ab eis differt folio asymmetrico, 
inflorescentia <$ valde ramosa multiflora, et cotyle- 
donibus multiplicatis. T: BW 7936. 

Arbor magna usque ad 45 m alt a et 1 m diam. 
Folia tenuiter coriacea, oblique ovato-elliptica, 
(5-)8-ll(-14) x (3-)4-6(-8) cm, ind. 1,5-2, glabra, 
nervis lateralibus uniparibus usque ad 4 / 5 partem 
laminae longitudinis ascendentibus, reticulatione 
laxa subscalariformi; petiolus applanatus vel ± 
sulcatus, 8-15 x 1-1,5 mm. Inflorescentiae <$ et $; 
<5 multiramosae, 60-150-florae; $ 5-10-florae. 
Fructus ovoideo-globosus, ± 4- vel 5-angulatus, 
10-12 x 8-10 mm; endospermium nullum; cotyle- 
dones multiplicati; embryo cur vat us. 

Large tree up to 45 m, 100 cm 0. Buttresses up 
to 2 3 / 4 m tall, 2 m out and 10 cm 0. Bark smooth 
to finely fissured, often pustullate, light-brown to 
grey-brown. Innovations densely rufous to yellow- 
ish-brown appressed-hairy, glabrescent. Terminal 
buds ovoid-conical, acute, c. 4-6 by 3 mm. 
Stipules ovate-acute, peltately attached and 
overlapping, thick, c. 5 by 3 mm. Leaves thin- 
coriaceous, obliquely ovate-elliptic, (5-)8-ll(-14) 
by (3-)4-6(-8) cm, index l'/ 2 -2; glabrous; often 
discolorous, upper surface dull grey-green, lower 
surface dull chocolate- or grey-brown; base 
attenuate-rounded, mostly asymmetrical; margin 
entire, often undulate; apex acute to acuminate; 
midrib and main lateral nerves raised beneath, 
impressed or flatfish above; nerves one pair, 
ascending and arcuating, running up to 4 / 5 or the 
entire length of the leaf; reticulations lax, sub- 
scalariform, rather distinct beneath; petiole 
glabrous, 8-15 by l-l'/jinm, flat, ! sulcate. 
Inflorescence <J or $, axillary or subterminal on 
new shoots, much-branched, many-flowered, 
including the bracts rather densely yellowish-brown 
to rufous appressed-hairy, glabrescent; bracts 
ovate-acute, c. 2 3 by I -1 '/a mm. flowers 5- 
merous. ', Inflorescences much-branched 

panicle, 60 I S()-llowered ; " (lower c. 2 mm 0; 
perianth lobes c. l'/j-2 by 1 mm; filaments l-l'/i 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 23. Lea, shape and venation of ™^^3?tc^^*^^^ 
Planch. — c. C. luzomca Warb. — d-e. C. tetranara koxb i u. hildebrandii Soepadmo 

Blume mi., / bb 33845,y Brass & Versteegh 11168, /c Lam 3635, / Brass z**w). 



1977] 



Ulmaceae (Soepadmo) 



65 



mm, anthers subreniform, c. 3 / 4 -l by 1 l 2 mm; 
pistillode ovoid-cylindrical, compressed, c. 3 / 4 by 
1 I 2 mm. — $ Inflorescence a much-branched 
raceme, up to 4-5 cm long, 5-10-flowered; 2 
flower ellipsoid, c. 3 by 2 mm ; perianth lobes ovate- 
acute, c. 2-3 by 1 mm; filaments l-lV 2 mm, an- 
thers c. 1 I 2 by 7a rnm; ovary ovoid-ellipsoid, c. 2 by 
1 mm, slightly compressed, initially densely hairy, 
glabrescent; stigmatic arms initially curved, later 
spreading, e. l-l^ram long, broadened and 
shallowly bilobed at the tip. Infructescence 3-5 cm 
long, axes glabrous, c. 1-2 mm 0, bearing 2-5 
fruits. Fruit ovoid-globose, glabrous, ± 4-5- 
angular, slightly beaked, 10-12 by 8-10 mm; 
exocarp thin, endocarp pitted; embryo curved, 
hypocotyle ascending, cotyledons broad, folia- 
ceous, folded; endosperm absent. 

Distr. Solomons (common); in Malesia: 
Moluccas (Buru, rare), New Guinea (W. & E. 
parts, common ; New Britain). 

Ecol. In both primary and secondary forests at 
0-1000 m; often very common and gregarious 
locally; on various types of soil. Male flowers 
appear with new shoots around Nov.-Dec., and 5 
ones around Jan. -Feb. ; fruits mature by May-June. 
The ripe fruits, which turn deep purple or bluish 
black, may be dispersed by birds. 

Vern. New Guinea: biheg, Hattam, walik, 
Mooi, sehiega, Manikiong, pi€h, Kebar, biempai, 
Biak; Solomons: laussi, lai-lai, Kwara. 

7. Celtis timorensis Span. Linnaea 15 (1841) 343; 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 315; Bl. 
Mus. Bot. 2 (1856) 71; Miq. Fl. Ind. Bat. 1, 2 
(1859) 22; Planch, in DC. Prod. 17 (1873) 180. — 
C. cinnamomea Lindl. ex Planch. Ann. Sc. Nat. 
Ill, 10 (1848) 303; Bl. Mus. Bot. 2 (1856) 72; 
Planch, in DC. Prod. 17 (1873) 181 ; Kurz, For. 
Fl. Burma 2 (1877) 472; Hook./ Fl. Br. Ind. 5 
(1888) 482; Prain, Beng. PI. 2 (1903) 719; Brandis, 
Ind. Trees (1906) 596; J. J. Smith in K. & V. Bijdr. 
12 (1910) 644; Merr. En. Philip. 2 (1923) 32; 
Gagnep. Fl. Gen. I.-C. 5 (1927) 682; Back. & 
Bakh./. Fl. Java 2 (1965) 1 1. — C. reticulosa Miq. 
PI. Jungh. (1851) 69; Fl. Ind. Bat. 1, 2 (1859) 222. 
— C. hamata Bl. Mus. Bot. 2 (1856) 72; Planch. 
in DC. Prod. 17 (1873) 180. — C. waitzii Bl. Mus. 
Bot. 2 (1856) 71 ; Miq. Fl. Ind. Bat. 1,2 (1859) 221 ; 
Planch, in DC. Prod. 17 (1873) 180. — C. dyso- 
doxylon Thw. En. PI. Zeyl. (1861) 267. — C. 
crenato-serrata Merr. Philip. J. Sc. 5 (1910) Bot. 
174. — Fig. 20f, 23f-h. 

Medium-sized tree up to 20 m, 30 cm 0. Bark 
smooth, grey. Innovations densely rufous-hairy. 
Branchlets glabrous, rather densely lenticellate. 
Terminal buds ovoid-conical, c. 3-4 by 2 mm. 
Stipules linear-lanceolate, 5-10 by 1-2 mm. Leaves 
thin-coriaceous, ovate-elliptic to elliptic-oblong, 
(6-)10-12(-17) by (2" 2 -)4-5(-8) cm, index |*/j 
2 l l 2 ; except for the midrib and nerves glabrous, 
discolorous, upper surface dull grey to blackish, 
lower surface chocolate-brown; base rounded to 
subcordate, mostly asymmetrical, margin undulate, 
distantly serrulate to crenate at least for the upper 
half; apex acute to acuminate; midrib and nerves 
flattish to impressed above, raised beneath; nerves 
I 2( J) pairs, arcuating and ascending, the lowest 
pair running to C/j-^/jf-Vj) the length of the leaf, 
anastomosing along the margin; reticulations 



subscalariform, sparse, rather distinct beneath; 
petiole 5-15 by 1-2 mm, sulcate. Flowers 5-merous. 
Inflorescence 3 or 6* 5, racemose, lax, axes slender, 
always longer than the petiole, including the bracts 
sparsely rufous-hairy; bracts narrow ovate-acute, 
c. 3-5 by 1-2 mm. — $ Inflorescences borne on the 
lower and leafless parts of the new shoots, much- 
branched, 10-20-flowered, up to 3 cm long; <S 
flower c. 2 mm ; perianth lobes c. 1 1 / 2 -2 by 1 mm ; 
filaments 1— l x / 2 mm, anthers reniform, c. 1 by 
7a mm; pistillode present but strongly reduced in 
size. — ^ Inflorescences borne in the axils of new 
leaves, 4-7-flowered, up to 2-3 cm long; $ flower 
c. 2-3 mm ; perianth lobes c. 1 1 / 2 -2 by 1 mm ; 
filaments c. 1 / 3 -l mm, anthers subreniform, 
c. ' 2 - 3 4 by l l 2 mm; ovary ovoid-ellipsoid, c. 2 by 
lmm; stigmatic arms c. 1-1\' 2 mm, spreading. 
Infructescence up to 4-5 cm long. Fruit ovoid, 
terete or 4-angular, beaked, 5-10 by 3-6 mm, 
glabrous. Embryo curved, hypocotyle ascending; 
cotyledons broad, folded, equal ; endosperm scanty. 

Distr. Ceylon, India, Bangladesh, Burma, 
Thailand, Indo-China; in Malesia: Central Suma- 
tra (rare), Java (common; incl. Christmas I.), 
Lesser Sunda Islands (Flores, Timor), N. Borneo 
(Mt Kinabalu), Philippines (Luzon). 

Ecol. Outside Malesia the species grows in 
evergreen forests or along streams in deciduous 
forests. In Malesia it is more common in areas 
with a rather prominent seasonal climate, 
0-1500 m. Fl.fr. Nov.-April. 

Vern. Central Sumatra: ki tondok, si tjakik; 
Java: jeungil, ki tamiang, S, tjengkek, tjitik, J; 
Philippines: takulao, Ibn., malabutulan, Tag. 

8. Celtis rubrovenia Elmer, Leafl. Philip. Bot. 2 
(1908) 464; Merr. En. Philip. 2 (1923) 33. — C. 
similis Merr. & Perry, J. Arn. Arb. 22 (1941) 253. 
- Fig. 23j. 

Deciduous tree up to 30 m, 60 cm 0. Bark grey- 
brown, smooth. Innovations densely rufous 
appressed-hairy, glabrescent. Older branchlets 
glabrous, sparsely lenticellate. Buds ovoid-conical, 
c . 3 by 2 mm, bracts ovate, imbricate, c. 1 by l j 2 mm. 
Stipules subulate, c. 3-4 by 1 mm. Leaves (fully 
developed ones) thick-coriaceous, rigid, brittle, 
ovate to ovate-elliptic, (3-)4-5(-6V 2 ) by (P/4-) 
1 l l 2 -2 l l 2 (-3 l l 2 ) cm, index 2-3.3 ; sparsely appressed- 
hairy especially on the midrib and nerves, glabres- 
cent; base attenuate-rounded to rounded, sym- 
metrical to asymmetrical; margin of young leaves 
serrate to crenate at least at the upper half, that of 
old leaves entire or distantly serrate in the upper 
half only; apex acute, acuminate to caudate, tip 
usually very sharp; midrib and nerves strongly 
raised beneath (reddish in fresh specimens), 
impressed above; nerves ( 1 — )2(— 3) pairs, arcuating, 
ascending, at a narrow angle with the midrib less 
than 45\ lowest pair running to c. 2 l 3 the length of 
the leaf; reticulations fine, subareolate, obscure on 
both surfaces; petiole (3-)4-6(-8) by 1 mm, 
deeply sulcate. Flowers 4-merous. Inflorescences 
6" or o*5, racemose, 1-2 cm long, 4-7-flowered. — 
[ Inflorescence borne on the lower and leafless 
parts of the new shoot, 5-7-flowered; (J flower 
c . 2 mm ; perianth lobes c. 1 i j 2 by I mm, ciliate ; 
filaments I I ' ', mm long, anthers reniform, 
c. I by J / 4 mm; pistillode minute. — Mixed (<$ $) 
inflorescences 4-5-flowercd, borne in the axil of 



66 



Flora Malesiana 



[ser. I, vol. 8 2 



new leaves; £ flower borne on the upper part of 
the inflorescence, c. 2-2 x j 2 mm ; perianth lobes 
c. I 1 1 2-2 by 1 mm, ciliate; stamens slightly smaller 
in size than those of the <$ flower; ovary ovoid, 
I 1 1 2-2 by 1 mm, densely hirsute; stigmatic arms 
spreading, lV2-2mm long. Infructescences up to 
lV 2 cm long, carrying (l-)2(-3) fruits. Fruit 
globose, c. 3 by 3 mm, glabrous, ± 4-angular. 
Endosperm scanty, oily, transparent. Embryo 
curved, hypocotyle ascending; cotyledons equal, 
foliaceous, folded. 

Distr. Malesia: Philippines (Luzon), New 
Guinea (Kebar Valley, Baliem R. ; Morobe Distr.). 

Ecol. In primary forests on hills, ridges between 
500-1800 m. In New Guinea sometimes rather 
common locally. Fl. fr. Sept.-March. 

Vern. Philippines: palek; W. New Guinea: 
dotjoni, nitjoni, Kebar. 

9. Celtis tetrandra Roxb. Fl. Ind. ed. Carey 2 
(1832) 63; Planch. Ann. Sc. Nat. Ill, 10 (1848) 
300; in DC. Prod. 17 (1873) 179; Kurz, For. Fl. 
Burma 2 (1877) 472; Gamble, Man. Ind. Timb. 
ed. 1 (1881) 344; Hook./. Fl. Br. Ind. 5 (1888) 482, 
ind. var. hamiltonii Hook./, et var. mollis (Planch.) 
Hook. / ; Prain, Beng. PI. 2 (1903) 719; Brandis, 
Ind. Trees (1906) 596; J. J. Smith in K. & V. 
Bijdr. 12 (1910) 641; Gagnep. Fl. Gen. I.-C. 5 
(1927) 681 ; Back. & Bakh./ Fl. Java 2 (1965) 11. 
— C. trinervia Roxb. Fl. Ind. ed. Carey 2 (1832) 65, 
non Lamk, 1797. — C. acata Hamilt. Trans. Linn. 
Soc. 17 (1834) 211; Planch. Ann. Sc. Nat. Ill, 
10 (1848) 299. — Sponia tetrandra (Roxb.) Voigt, 
Hort. Suburb. Calc. (1845) 294. — C. napalensis 
Planch. Ann. Sc. Nat. Ill, 10 (1848) 298. — C. 
glabra Planch. I.e. 298. — C. roxburghii Planch. 
I.e. 302. -— C. hamiltonii Planch. I.e. 301 ; in DC. 
Prod. 17 (1873) 179. — C. mollis Planch. Ann. Sc. 
Nat. Ill, 10 (1848) 297, p.p., quoad specim. ex 
Wallich 7203; in DC. Prod. 17 (1873) 179. — C. 
serotina Planch. Ann. Sc. Nat. Ill, 10 (1848) 
301 ; Wight, Ic. 4, 4 (1850) t. 1570. —Fig. 181-o, 
23d-e. 

Semi-deciduous tree up to 40 m and 100 cm 0. 
Bark smooth to rough, grey-brown. Innovations 
densely rufous-hairy, glabrescent. Terminal bud 
ovoid-globose, 2-3 mm 0. Stipules linear, c. 5 by 
1 mm. Leaves coriaceous, oblique ovate-elliptic, 
(4-)6-10(-13) by (lVa-^ 1 ^ 1 ^) cm, index 2-3, 
broadest at or below the middle; slightly dis- 
colorous, above glabrous, dull grey in drying, 
beneath glabrous or sparsely rufous-pubescent 
especially on midrib and nerves, yellow grey-green 
in drying; base rounded-attenuate, strongly asym- 



metrical; margin denticulate or distinctly serrate 
at the upper half, or subentire ; apex acute, acumi- 
nate, to caudate; midrib and nerves flat to im- 
pressed above, raised beneath; nerves 3-4 pairs, 
ascending and arcuating, anastomosing near the 
margin, the lowest pair running to c. 1 l 3 - 2 / 3 the 
length of the leaf; reticulations fine, subscalariform, 
rather distinct beneath; petiole 5-12 by 1-2 mm; 
slightly sulcate. Inflorescence <$ or $, cymoid, in- 
cluding the bracts densely rufous-hairy. — S In- 
florescence borne on the lower and leafless part of 
the new shoot, 2-5-flowered, occasionally several 
of them are clustered together on leafless short 
lateral new shoots giving rise to a raceme of cymes ; 
3 flower c. 2-3 mm ; perianth lobes 4(-5), 
c. I 1 1 2-2 by 1 mm, ciliate; filaments V2-I mm, 
anthers reniform, c. 1 by V2 mm, sparsely seri- 
ceous; pistillode very much reduced. — $ 
Inflorescences 2-5-flowered, axes slender, in the 
axils of new leaves; £ flower c. 2 mm ; perianth 
lobes ciliate, c. 2 by 1 mm; stamens as in the S 
flower; ovary ovoid, c. 2 by 1 mm, sparsely minute 
pubescent; stigmatic arms c. 2 mm long, 1 j 2 mm 
broad, spreading. Fruit ± globose, 5-8 mm 0, 
glabrous, turning deep-red or black when ripe. 
Endosperm scanty. 

Distr. India, Bangladesh, Burma, Thailand, 
Indo-China; in Malesia: N. Sumatra (Gajo and 
Karo Lands) Java (West : G. Galunggung ; Central : 
G. Muria; East: Bodjonegoro, Mts Kawi & Idjen, 
Besuki, etc.), Lesser Sunda Islands (Bali, Sumbawa, 
Flores). 

Ecol. In primary and secondary forests, 0-2000 
m. In Burma and Thailand very often in evergreen 
or semi-deciduous forest along river-banks. In 
Malesia the preference seems to be largely to areas 
subject to a seasonal climate. Fl.fr. Aug.-April. 

Vern. Sumatra: ndokum, sigar, Karo, bitatar, 
Toba, temung, M; W. Java: ki djeungkil, ki 
tamiang, S, tiritih, tritih, J ; Lesser Sunda Is. : pusu, 
Sumbawa, namut, Flores. 



Excluded 

Celtis grewioides Warb. Bot. Jahrb. 13 (1891) 287 
= Ziziphus angustifolius (Miq.) Hatus. Nova 
Guinea Bot. n. 3 (1960) 13. 

Solenostigma angustifolium Miq. Sumatra (1861) 
412. — C. angustifolia (Miq.) Planch, in DC. 
Prod. 17 (1873) 186, non Lindl. ex Wall. Cat. 
(1831) n. 3691, nomen = Ziziphus angustifolius 
(Miq.) Hatus. Nova Guinea Bot. n. 3 (I960) 13. 



5. APHANANTHE, mm. gen. cons. 

Planch. Ann. Sc. Nat. Ill, 10 (1848) 265, non Link, 1821 ; Miq. Fl. Ind. Bat. 1, 2 
(1859) 218; Planch, in DC. Prod. 17 (1873) 207; Hook./, in B. & H. Gen. PI. 3 
(1880) 355; Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 66; Bernard, Bull. Herb. 
Boiss. II, 6 (1906) 34; Gagnep. Fl. Gen. I.-C. 5 (1927) 690; Leroy, Bull. Mus. 
Hist. Nat. Paris II, 18 (1946) 118, 180; Fl. Madag. Fam. 54 (1952) 12; J. Agr. 
Trop. Bot. Appl. 8 (1961) 72; Li, Woody Fl. Taiwan (1963) 105; Hutch. Gen. Fl. 



1977] Ulmaceae (Soepadmo) 67 

PL 2 (1967) 149. — Homoioceltis Bl. Mus. Bot. 2 (1856) 64. — Galumpita Bl. I.e. 
73; Miq. Fl. Ind. Bat. 1, 2 (1859) 223. — Gironniera subg. Galumpita Hook./, in 
B. & H. Gen. PI. 3 (1880) 356; Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 66. — 
Mirandaceltis A. J. Sharp, Bol. Soc. Bot. Mex. 23 (1958) 38, f. 1-4. — Fig. 24. 

Monoecious, deciduous or semideciduous shrubs or trees, often buttressed. 
Innovations densely or sparsely, whitish-grey to rufous, appressed-pubescent. 
Indumentum consisting of bulbous-based, unicellular, finely tuberculate hairs and 
multicellular, glandular hairs. Older branches glabrous, lenticellate, bearing lateral 
and terminal buds. Stipules lateral, extrapetiolar, subulate, caducous. Leaves 
alternate, petioled, glabrous, coriaceous, triplinerved at base or pinnately nerved. 
Inflorescences <J, $, or very rarely <£$, axillary; bracts minute, caducous. — 6* 
Inflorescences a condensed, multi-flowered raceme, borne on the lower parts of the 
new shoots; <$ flowers short-stalked, globular, 4-5-merous; perianth lobes mem- 
branous, imbricate in bud, sparsely appressed pubescent outside ; stamens glabrous, 
filaments subulate, inflexed in bud, anthers ovoid-subreniform, non-apiculate, 
introrse ; pistillode absent, replaced by a cluster of whitish to silvery, erect, soft, 
simple hairs. — $ Flowers solitary in the axil of the upper leaves of the new shoot, 
or borne in a 2-3-flowered mixed (<^$) racemose inflorescence; long-stalked; 
perianth lobes 4-5, long-persistent; staminode absent; ovary sessile, ovoid-ellip- 
soid, terete to angular; stigmatic arms tubular; ovule anatropous. Drupe fleshy, 
ovoid-globose, faintly 3-5-angular or ± terete, glabrous; endocarp hard and per- 
sistent. Seed exalbuminous, coat membranous, few cells thick; embryo curved, 
hypocotyle ascending, cotyledons more or less equal, involute. Mode of germina- 
tion unknown. 

Distr. About 4-5 spp. Mexico, Madagascar, Ceylon, India, Burma, China (also Hainan), Korea, 
Japan, Taiwan, Hongkong, Indo-China, Thailand, Andamans, through Malesia to Australia (Queensland 
and New South Wales) and Solomons. Throughout Malesia (except Malaya and Moluccas) : 2 spp. Fig. 25. 

Ecol. In Malesia mainly found in areas subject to a rather strong seasonal climate, on various types of 
soil in the coastal lowlands, hills, and gallery forests, 0-750 m, locally often abundant and forming dense 
thickets. 

In the north temperate and subtropical regions the species flower in April-May and drupes ripen in 
July-August. Tropical species produce flowers twice a year, viz around March-April and Sept. -Oct. and 
fruit ripens in June-July or Nov.-Dec. 

The deciduous or semideciduous habit, flush-wise mode of growth, structure, size, colour of the in- 
florescence and flowers suggest that pollination is affected by wind. The drupes which turn to a deep red 
colour when ripe are possibly dispersed by frugivorous birds. 

KEY TO THE SPECIES 

I. Leaves ovate, elliptic or obovate, (2-)4-6(-10) by (l-)2-3(-4)cm, triplinerved at base, margin variously 
serrate or dentate, very rarely subentire; nerves (3-)4-5(-6) pairs, at a 30-45° angle with the midrib. 
Mature fruit 6-10 by 4-7 mm, beak 1-3 mm 1. A. philippinensis 

I. Leaves ovate-elliptic or elliptic-oblong, (5-)10-14(-20) by (2-)3-6 (-8) cm, pinnately nerved, margin 
entire or very rarely distantly, minute serrulate at the upper half; nerves (5-)7-8(-10) pairs, at an angle 
of more than 60 with the midrib. Mature fruit 15-20 by 8-12 mm, beak up to 5 mm. . 2. A. cuspidata 

1. Aphananthe philippinensis Planch. Ann.Sc. Nat. Shrub to medium-sized tree up to 28 m, 40 cm 0. 

Ill, 10 (1848) 337; Miq. Fl. Ind. Bat. I, 2 (1859) Trunk often fluted, low-buttressed, occasionally 

219; Planch, in DC. Prod. 17 (1873) 208; Bth. producing suckers. Bark smooth to finely fissured, 

Fl. Austr. 6 .1873) 160; Hook./. Icon. Ill, 2 (1876) peeling off into rectangular flakes, lenticellate. 

65, t. 1741 ; Mkrr. En. Philip. 2(1923) 34; Fkancis, Young parts densely or sparsely greyish-brown or 

Austr. Rain-forest Trees ed. 2 (1951 ) f. 24 <Sc 25. rufous, short, simple pubescent. Stipules subulate, 

— Taxotrophis rectinervia F.v.M. Fragm. 6 (1863) 2 3 by ',', mm. Leaves glabrous, thin- to thick- 

192. A m imrrvui (F.V.M.) PLANCH, in DC. coriaceous, ovate, elliptic, or obovate, (2 )3 6( 10) 

Prod. 17 (1873) 208. — Fig. 24j n. by (l-)2-3(-4) cm, index 2-2.6; base attenuate 



68 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 24. Aphananthe cuspidata (Bl.) Planch, a. Habit, nat. size, b. LS of mature fruit, x 1V 3 , c. basal 
view of fruit, nat. size, d. flowering young shoot, nat. size, e. 6* flower before anthesis, x 6, f. ditto in 
section, x 9, g. cJ flower at anthesis, x 13, h. $ flower, x 8, i. ditto in section, x 16. — A. philippinensis 
Planch, j. mature fruit, x 2, k. ditto, basal view, x l 2 / 3 , 1-n. variation of leaf-form, x 2 / 3 (a-c Koorders 
21330, d-i Koorders 30071, j, k, m, n Borden FB 1286, / Ramos BS 27383). 



1977] 



Ulmaceae (Soepadmo) 



69 




Fig. 25. Range of Aphananthe Planch. — A. cuspidata (Bl.) Planch, (dots), A. philippinensis Planch. 
(stars), A. sakalava Leroy (square), A. aspera Engl, (circles), A. monoica (Hemsl.) Leroy (triangles). 



or rounded, more or less equal-sided; margin 
serrate, dentate, or rarely subentire, ends of serra- 
tion occasionally developing into sharply mucro- 
nate structures; apex acute, or rounded, tip blunt; 
midrib and nerves strongly raised beneath, 
flattish above; nerves (3-)4-5(-7) pairs, ascending, 
straight or arcuating, subparallel, at 30-45° with 
the midrib, not anastomosing near the margin; 
reticulations fine, lax, subscalariform, faintly 
visible beneath; petiole (2-)3-5(-7) by 1 mm, 
adaxially flat or shallowly sulcate. — <$ Inflores- 
cence 1-3 cm 0, 10-50-flowered; bracts linear- 
acute or narrow ovate-acute, V 2 -l by V^mm; rj 
flowers 1-1' 2 mm 0; perianth lobes ovate-lanceo- 
late, 1-1' 2 by ' 2 1 mm; filaments I / 2 - 3 / 4 mm, 
anthers c. 3 4 by ' 2 mm. — 5 Flowers always 
solitary, ovoid-ellipsoid, 1-1' 2 by 1 mm, c. 5 mm 
pedicelled; perianth lobes narrow ovate-acute, 
1 2-1 by ' 2 mm; ovary I— l l /a by 1 mm, densely 
appressed-hairy ; stigmatic arms 2-3 mm. Fruit 
ovoid-globose, 6-8 by 4-6 mm, 3-4-angular, 
sparsely appressed-pubescent, glabrescent, pedicel 
c. 5-10 mm. 

Distr. Australia (Queensland and New South 
Wales) and Solomons; in Malesia: New Guinea 
and Philippines (Luzon). Fig. 25. 

Ecol. In primary and secondary forest subject to 
a rather strong seasonal climate, 0-750 m. In New 
Guinea it is often found in semi-deciduous 
gallery- or mixed Eucalyptus-fontt, occasionally 
forming dense thickets especially on hillsides. Ft. 
Apr. May and Sept. Oct., fr. mature in July Aug. 
and Nov. Dec. 

2. Aphananthe cuspidata (Mi.) PLANCH, in DC. 
Prod. 17 (1873) 20'>. Cychstemon cuspidatum 
Hi . Bijdr. (1825) 599. GaJumpita cuspidata (Bl.) 

Bl. Mus. Hot. 2 (1856) 73; Mio. H. Ind. Bat. I, 2 



(1859) 224. — Gironniera nitida Bth. Fl. Hongk. 
(1861) 324. — Gironniera reticulata Thw. En. PI. 
Zeyl. 1 (1861) 268; Hook./. Fl. Br. Ind. 5 (1888) 
486. — Gironniera lucida Kurz, For. Fl. Burma 2 
(1877) 470; Hook./. Fl. Br. Ind. 5 (1888) 486. — 
Gironniera cuspidata (Bl.) Kurz, For. Fl. Burma 2 
(1877) 470; J. J. Smith in K. & V. Bijdr. 12 (1910) 
688; Merr. En. Philip. 2 (1923) 35; Back. & 
Bakh. /. Fl. Java 2 (1965) 12. — Gironniera 
curranii Merr. Philip. J. Sc. 4 (1909) Bot. 251. — 
A. lissophylla Gagnep. Bull. Soc. Bot. Fr. 72 (1925) 
804; Fl. Gen. I.-C. 5 (1927) 690. — Gironniera 
thompsoni King ex A. M. & J. M. Cowan, Trees 
N. Bengal (1929) 122. — ^Gironniera yunnanensis 
Hu, Bull. Fan Mem. Inst. Biol. Bot. Ser. 10 (1940) 
150. — 1A. yunnanensis (Hu) Grudz. Nov. Syst. 
PI. Vase. USSR (1964) 66. — Fig. 24a-i. 

Small to medium-sized tree up to 30 m, 60 cm 0. 
Buttresses up to 1 m tall, and 2 m out. Bark surface 
rough, grey-brown, often flaky. Young branchlets 
sparsely, minutely, appressed-pubescent, glab- 
rescent; older branchlets sparsely lenticellate. 
Stipules narrow ovate-acute, 2-3 by 1 mm. Leaves 
coriaceous, glabrous, ovate-elliptic to elliptic- 
oblong, (5-)10-14(-20) by (2-)3-6(-8) cm, index 
2'/ 2 -3; base rounded, subcordate, or attenuate, 
equal-sided or occasionally slightly unequal; 
margin entire, rarely distantly, minute serrulate in 
the upper half, often undulate; apex acute, acumi- 
nate, or cuspidate, acumen up to 2 cm, sharp- 
tippcd; midrib strongly raised and prominent 
beneath, impressed to llattish above; nerves (5 )7 
Hi 10) pairs, slightly raised beneath, flattish above, 
subparallel, arcuating, at an angle of more than 
60 with the midrib, weakly anastomosing near the 
margin; reticulations line, indistinct on both 
lUrfaCM, lubtcalariform to sub-arcolate; petiole- 
glabrous, (5 )8 I2( 15) by 1-2 mm, sulcate. — <$ 



70 Flora Malesiana [ser. I, vol. 8 2 

Inflorescence up to 4 cm, 10-30-flowered; bracts land forests subject to a rather pronounced seasonal 

ovate-acute, V4-V2 DV V4 mm; <J flower \ l l 1 -2 mm climate. In Thailand it occurs mainly in the ever- 

0, short-stalked; perianth lobes obovate-lanceo- green or semi-deciduous forests along streams, 

late, c. 2 by 1 mm; filaments l-l l l 2 mm, anthers Outside Malesia flowering in March-April and 

ovoid-subreniform, c. 1 mm 0. — 9 Flower fruits mature in July-August. In Malesia it flowers 

solitary or borne in a 2-3-flowered mixed inflores- twice yearly, viz in Febr. -March and Sept.-Oct., 

cence, ovoid-ellipsoid, c. 2 by 1 mm; pedicels up to fruits maturing in June-July and Nov. -Dec. 

10mm; perianth lobes coriaceous, ovate-acute, Vern. Java: suluh, wuluh; Lesser Sunda Is.: 

c. 2 by 1 mm; ovary ovoid, glabrous, c. IV2 by sulu, Bali, k. belikat, Sumbawa, k. loko, Flores. 

lmm; stigmatic arms 2-3 mm. Fruit ovoid, Note. Though I have not been able to examine 

glabrous, including the beak 15-20 by 8-12 mm, any of the specimens cited by Hu (1940), judging 

beak up to 5 mm; pedicel up to 3 cm, 1 mm 0. from the description given it is likely that Giron- 

Distr. Ceylon, India, Burma, Andaman Is., ? niera yunnanensis Hu must be referred to this 

China, Hainan, Hongkong, Indo-China, Thailand; species, 
in Malesia: NE. Sumatra (Sibolangit, doubtfully 

native), Java (mainly N. Central & E. parts), Excluded 
Lesser Sunda Islands (Bali, Sumbawa, Flores, 

Timor, rather common), Borneo (very rare, Aphananthe negrosensis Elmer, Leafl. Philip. 

Kinabalu area), Philippines (Mindanao, rare), Bot. 2 (1909) 575. — Calaunia negrosensis (Elmer) 

Celebes (rare). Fig. 25. Grudz. Nov. Syst. PI. Vase. USSR (1964) 54 = 

Ecol. In primary lowland to submontane forest, Streblus glaber (Merr.) Corner, Gard. Bull. 

0-1300 m. In Malesia it is mainly confined to low- Sing. 19 (1962) 221 (Moraceae). 

6. GIRONNIERA 

Gaudich. Voy. Bonite (1844) t. 85; Planch. Ann. Sc. Nat. Ill, 10 (1848) 338; 
Bl. Mus. Bot. 2 (1856) 72; Miq. Fl. Ind. Bat. 1, 2 (1859) 222; Planch, in DC. 
Prod. 17 (1873) 205; Hook./, in B. & H. Gen. PI. 3 (1880) 356; Fl. Br. Ind. 5 (1888) 
485; Engl, in E. & P. Nat. Pfl. Fam. 3, 1 (1888) 66; Bernard, Bull. Herb. Boiss. II, 
6 (1906) 34, map 24; J. J. Smith in K. & V. Bijdr. 12 (1910) 665; Hutch. Gen. Fl. 
PL 2 (1967) 149; Phuphathanaphong, Thai For. Bull. 6 (1972) 49; Soepadmo in 
Whitmore, Tree Fl. Mai. 2 (1973) 417. — Nemostigma Planch. Ann. Sc. Nat. Ill, 
10 (1848) 265, cf. p. 338. — Helminthospermum Thw. in Hook. J. Bot. Kew Misc. 6 
(1854) 301, t. 9c. — Fig. 26. 

Shrubs or large-sized trees, very rarely buttressed. Bark smooth to finely fissured, 
grey-brown, often lenticellate. Innovations densely or sparsely set with golden 
yellow or yellowish-brown indumentum consisting of simple bulbous-based, finely 
tuberculate, appressed or erect hairs and multicellular capitate-glandular hairs. 
Stipules extrapetiolar, free but overlapping each other and completely enclosing 
the bud, on falling leaving a circular scar around the node. Leaves pinnately 
nerved, nerves parallel, regularly well-spaced. Inflorescence <$ or $, very rarely 6*$, 
axillary or borne on older, leafless branchlets, 1 -many-flowered, paniculate, race- 
mose, thyrsoid, or capitate ; bracts ovate-acute, minute, caducous, but rather long 
persistent in the $ inflorescence. — <$ Flower globular, sessile or short-stalked along 
the axes; perianth lobes 5, imbricate in bud; stamens glabrous, anthers introrse; 
filament subulate, inflexed; anthers ovoid-reniform, apiculate, sub-basifixed ; 
pistillode present, rather well-developed or very rudimentary, densely set with 
whitish or silky erect hairs. — $ Flower ovoid-ellipsoid, compressed ; perianth lobes 
4-5, usually unequal in size, long persistent; staminodes absent; ovary ovoid- 
ellipsoid, strongly compressed, sessile, densely or sparsely appressed-hairy, 
glabrescent ; stigmatic arms up to 1 1 / 2 cm long, curled in bud, later spreading, rather 
long persistent; ovule anatropous. Drupe ovoid-globose, convex elliptic lens- 
shaped; exocarp thin, strongly adnate to the hard and persistent endocarp. Seed 



1977] 



Ulmaceae (Soepadmo) 



71 




Fig. 26. (jtronmera celtidifolia Gaudich. a-b. V Flowers, one in CS, c-g. <J flowers, various details — 
G. subaequalis Pi anc h. h. J Mower, m. young infructescence, p-r. fruits, one in LS. — G. nervosa 
Flam h i j | flowers. O. parvifolia Planch, k. f Inflorescence, n. young infructescence — G 
rnamnifoliu Hi I J Inflorescence. C. /uWu Rim . o. Young infructescence. a, c, m r X J /, A • 3 </-<>• 

* v\l!u< flU> ! ^r' , 2 ;* ' '' ,( " ''^^^'.^MauwanaoII^ARahmatsi'boha^II; 
lyKLPIRl 14582, k KLP PRI 18017, / Zipph ins s.n.. m Mom. Shah 519, n SF 34908. o bb 30873 p r 

K f-.P/FRI 99766). 



72 



Flora Malesiana 



[ser. I, vol. 8 2 



coat membranous, a few cells thick; endosperm absent; embryo curved, hypo- 
cotyle ascending; cotyledons fleshy, narrow, equal. Mode of germination unknown. 

Distr. 6 spp., SE. Asia: Ceylon, Andaman Is., Burma, Thailand, Indo-China, China (Yunnan, 
Kwangtung, Canton, Hainan, Hongkong), Micronesia (Palau and Ponape Is.), Melanesia (Solomons), 
Polynesia (Samoa, Fiji); in Malesia: throughout the region except for the eastern part of Java, the Lesser 
Sunda Islands, and the SE. Moluccas. Fig. 27. 




Fig. 27. Approximate range of the genus Gironniera Gaudich. with the number of species in each island or 

partial area. There are no endemic species. 



Fossils. Wolfe (in Graham, Flor. & Paleoflor. Asia & N.E. America, 1972, 200, pi. IV, f. 2) reported 
leaf-impressions attributed to Gironniera from the Early Ravenian Flora of Alaska (Middle Eocene). 
However, in examining the published photograph (p. 21 1), I am very much in doubt whether the specimen 
belongs to the genus at all. 

Ecol. Understorey shrubs or trees in both primary and secondary forests, at 0-1300 m, often abundant 
and gregarious; on various types of soil, including those derived from limestone. In areas where the 
seasonal climate is prominent mainly found in the evergreen forest along streams. 

Judging from the structure of the flower and inflorescence, it is inferred that the pollination may be 
affected by wind. The drupes which turn to bright yellow or orange in colour when ripe are mostly barren, 
and they may be dispersed by various species of frugivorous birds. Direct observations in the field are, 
however, wanting. 

Notes. Gironniera spp. have a continuous, flush-wise growth habit and have the ability to produce 
flowers and fruits at a very young (sapling) stage (2-3 m tall). Since the plants are often very common 
locally both in the primary and secondary forests, produce flowers and fruits regularly, and are very easy 
to collect, most of the examined specimens were gathered from these young plants. The presence of so 
many specimens collected from juvenile plants hampers proper identification even when they are fertile. 

In the present revision, the characters used in the key were taken from specimens collected from mature 
or fully grown trees, while those mentioned in the description of each species include also data from speci- 
mens collected from the young plants, thus to include the total morphological variability. 

On the material and field notes so far available it is impossible at this stage to determine whether the 
genus is strictly monoecious or dioecious. In most cases, the specimens display only fruits or $ inflores- 
cences, thus giving the impression that the genus is dioecious. However, there are a few collections (in all 
species but G. hirta) which have both <$ influorescences and infructescences attached to the same branch- 
let, or they are borne on separate branchlets belonging to a single collecting number. 

Mrs. Phuphathanaphong I.e. accommodated the Malesian specimens into two species, G. nervosa and 
G. celtidifolia, without argumentation. I cannot agree with this view. 



1977] Ulmaceae (Soepadmo) 73 

KEY TO THE SPECIES 

(Based on specimens from mature trees) 

1 . <J & 9 Inflorescence and infructescence a condensed, capitate thyrse or panicle, axillary or borne on 
older leafless branchlets. o Flower with a rather well-developed pistillode. Leaves thick-leathery, 

distantly serrate or subentire. Terminal bud (stipules) up to 4 1 / 2 by i l 2 cm 1. G. celtidifolia 

1. 3 & 9 Inflorescence and infructescence a lax, branched or unbranched panicle, raceme, or thyrse, 
always axillary. 5 Flower with a strongly rudimentary pistillode. Leaves chartaceous to thick leathery, 
in Mai. specimens mostly entire. Terminal bud less than 3 cm by less than 0.3 cm. 
2. Mature leaf densely pubescent beneath. £ Inflorescence a much-branched thyrse; 3 flowers arranged 
in condensed cymoid clusters of 5-10 along the axes. 
3. Young branchlets densely short appressed-hairy. Leaves elliptic-lanceolate to elliptic-oblong, index 
2V2-3, broadest at the middle; nerves (12— )15(— 17) pairs, parallel, straight, 5-8 mm spaced, not or 
only weakly anastomosing along the margin. Well-developed 9 inflorescence (thus also the infructes- 
cence) mostly an unbranched panicle, c. 3 cm long, bearing less than 10 flowers (fruits). 

2. G. nervosa 

3. Young branchlets densely set with long, erect hairs. Leaves mostly ovate-elliptic, rarely elliptic- 
oblong, index c. 2, broadest below or rarely at the middle; nerves (8-)10-12(-14) pairs, subparallel, 
10-15 mm spaced, arcuating and anastomosing towards the margin. Well-developed 9 inflorescence 
(cq. infructescence) a mostly much-branched panicle 5-10 cm long, bearing up to 20 fruits 

3. G. hirta 
2. Mature leaf glabrous. 6* Inflorescence unbranched or a branched raceme or panicle; 6* flowers 
solitary along the axes. 

4. Mature leaf thick-coriaceous, broad-ovate, elliptic, or elliptic-oblong, broadest below or at the 
middle, index 2-2\' 2 , base unequal-sided. Terminal bud 2-3 by 0.2-0.4 cm. Well-developed 3 
inflorescence a much-branched panicle, carrying 40-100 flowers. Well-developed 9 inflorescence 
(cq. infructescence) carrying (2-)5-10(-15) flowers (fruits). Mature fruits 10-12 by 9 by 5 mm. 

4. G. subaequalis 

4. Mature leaf chartaceous to thin-coriaceous, elliptic-lanceolate or elliptic-obovate, broadest at or 

slightly above the middle, index (2\ 2 -)3(-3\ 2 ), base more or less equal-sided. Terminal bud 2 by 

1 I J cm. Well-developed 3 inflorescence an unbranched panicle carrying 1 5-30 flowers. Well-developed 

9 inflorescence (cq. infructescence) carrying l-3(-5) flowers (fruits). Mature fruits 7-8 by 5-6 by 

3-4 mm. 

5. Nerves (5-)6-8(-9) pairs. S Flowers short-stalked. Axes of 9 inflorescence (infructescence) slender 

and thin (less than 0.3 mm), 4-10 cm long, carrying l-3(-5) flowers (fruits). . . 5. G. parvifolia 

5. Nerves 10-12 pairs. 6" Flowers sessile along the axes. Axes of 9 inflorescence (infructescence) 

V 2 -l mm thick, 3-4 cm long, carrying (l-)3-5 flowers (fruits) 6. G. rhamnifolia 

1. Gironniera celtidifolia Gaudich. Voy. Bonite terete or adaxially flat near the base, strigose, 

(1844) t. 85; Planch. Ann. Sc. Nat. Ill, 10 (1848) glabrescent. Inflorescences a condensed, capitate, 

340; Miq. Fl. Ind. Bat. 1, 2 (1859) 223; Seem, much-branched panicle or thyrse, axillary or borne 

Fl. Vit. (1865) 236; Planch, in DC. Prod. 17 on older, leafless branchlets, many-flowered, 6* or 

(1873) 207; Laut. Bot. Jahrb. 50 (1913) 326; 9, very rarely ^9, including the bracts densely 

Merr. En. Philip. 2 (1923) 35; Parham, PI. Fiji appressed-hairy; bracts 2-4 by 2 mm. — 6* Flower 

Is. (1964) 88. — G. sibuyanensis Elmer, Leafl. c. 17 2 -2mm 0, sessile or short-stalked along the 

Philip. Bot. 5 (1913) 1845; Merr. En. Philip. 2 axes, perianth lobes ovate-elliptic, concave, 

(1923) 35. — G. grandifolia Merr. & Perry, c. lV 2 -2 by l-l'/jrnm; filaments l'/ 2 -2mm, 

J. Arn. Arb. 20 (1939) 325. — G. retinervia Merr. anthers 1-1 V2 by 1 mm; pistillode cylindrical, 

& Perry, I.e. 326. — Fig. 26a-g. IV2-2 by l /a mrn > at base surrounded by dense, 

Shrub or medium-sized tree up to 18 m tall and whitish, erect hairs. — 9 Flower 2-3 by 2 mm; 

25 cm 0. Bark greyish-brown to dark-brown, perianth lobes ovate, I-IV2 by V2-I mm; ovary 

smooth or finely fissured. Terminal buds c. 4-4'/2 2-3 by l'/ 2 -2mm, densely appressed-hairy; 

by \ 2 cm; stipules narrow lanceolate-acute, stigmatic arms 1-1 '/ 2 cm, slender. Fruit ovoid-.. 

c. 4-4'/ 2 by l U- l l 2 cm. Leaves thick-coriaceous, ellipsoid, sparsely appressed-hairy, glabrescent, 

elliptic-, lanceolate-, or obovate-oblong, (8-)15- 5-8 by 4-6 by 3-4 mm, beak up to 5 mm. 
25M2) by (4-)6-12(-19) cm, index 2V2-372, Distr. Micronesia (Palau, Ponape), Melanesia 

broadest at or above the middle; base unequal- (Solomons), Polynesia (Samoa, Fiji); in Malesia: 

sided, attenuate, rounded, or cordate-auriculate; Borneo (doubtful record), Philippines (common 

margin distantly serrate at least for the upper half and widespread). Moluccas (Morotai, Halma- 

or subentire; apex rounded, acute, or acuminate- heira). New Guinea (common in W. & E. ; also in 

caudate, acumen up to 4 cm long; glabrous above, Admiralty, Misima, Sudest, and Rossel [&.). 
sparsely apprcsscd-hairy beneath especially on Ecol. In both primary and secondary forests, 

midrib and nerves, glabrescent; midrib and nerves 1200 in; often locally abundant as undcrstorey 

strongly raised beneath, flattish to shallowly im- shruh or tree. Fl. fr. Jan. Dec. 
pressed above; nerves (K )I2 IM 19) pairs. Vein. Philippines: oblattg, Many., iliiu, 1 1| ., 

anastomosing along the margin; reticulations matlgabau, His., taitglffttungan, Mho.; Moluccas: 

subscalariform, evenly spaced, distinct beneath; koko, MOFOtaij New Guinea: aimarwlrteh, Biak, 

petiole (3 )5 I2( 15) by (I )\' 2 2' 2 ( 4) mm, taun, Wapil tang., Sepik Distr.; Solomons: 



74 



Flora Malesiana 



[ser. I, vol. 8 2 



aisulia, Kwara; Fiji: tnasivau, sisisi; Samoa: 
pua-pua, puluvao. 

Notes. In most specimens examined, the 6* 
inflorescence shows abnormal development and 
produces a malformed structure consisting of 
numerous superimposed bracts in the axils of 
which are found sterile structures resembling in 
size and shape that of the pistillode in the normally 
developed and functionally 6* flowers. Well- 
developed and functionally S flowers are to be 
found only at the distal parts of these abnormally 
developed inflorescences. In a cJ9 inflorescence, 
which is very rarely present, the functionally $ 
flowers are borne on the central main axis and they 
are flanked by two or more lateral, functionally $ 
flowers. Most of the fruits so far examined (more 
than 95 %) are barren. 

Size and shape of the leaf are rather variable. In 
general, specimens collected from a higher altitude 
or from a more exposed habitat have a smaller 
leaf and less pairs of nerves than those gathered 
from lowland and shady localities. Intermediates 
are, however, not uncommon, and for this reason 
G. sibuyanensis, G. grandifolia, and G. retinervia 
are here reduced. 

2. Gironniera nervosa Planch. Ann. Sc. Nat. Ill 
10 (1848) 338; Bl. Mus. Bot. 2 (1856) 74; Miq 
Fl. Ind. Bat. 1, 2 (1859) 222; Planch, in DC 
Prod. 17 (1873) 206; Kurz, For. Fl. Burma 2 
(1877) 469; Hook. /. Fl. Br. Ind. 5 (1888) 485 
Merr. En. Born. (1921) 216; Ridl. Fl. Mai. Pen. 3 
(1924) 320; Corner, Ways. Trees (1940) 688 
Soepadmo in Whitmore, Tree Fl. Mai. 2 (1973) 
419, f. 2. — G. penangiana Gandog. Bull. Soc 
Bot. Fr. 66 (1919) 289. — G. sponioides Gandog 
I.e. 289. — Fig. 26i-j. 

Small to large-sized tree up to 40 m, 60 cm 0. 
Buttresses sometimes present, low. Bark smooth or 
finely fissured, grey-green to dark grey-brown, 
often hoop-marked and lenticellate. Terminal bud 
1-2 by 7 4 -7 2 cm; stipules 17 2 -2 by l / 4 -7 2 cm. 
Leaves thick-coriaceous, rigid, elliptic-lanceolate 
to elliptic-oblong, (6V2-)10-15(-18) by (2V 2 -)4-6 
(-8) cm, broadest at the middle, index 272-3 ; 
except for the midrib and lateral nerves glabrous 
above, densely set with yellowish-brown, soft, 
slender hairs beneath ; base rounded or attenuate, 
unequal-sided; margin entire, often recurved; apex 
rounded to acute; midrib and nerves strongly 
raised beneath, flatfish or impressed above ; nerves 
(12-)14-16(-17) pairs, 5-8 mm spaced, arcuating 
but not anastomosing near the margin, forming an 
angle of up to 60° with the midrib; reticulations 
dense, regularly spaced, scalariform or subscalari- 
form, strongly raised and prominent beneath, 
obscure to faintly visible above; petiole 5-10 by 
1-2 mm, subterete, densely yellowish-brown ap- 
pressed, pubescent. Inflorescences 6* or $, axillary, 
borne on separate shoots, including the bracts 
densely yellowish-brown, appressed-pubescent ; 
bracts narrow ovate-acute, c. 1-2 by 1 mm. — <$ 
Inflorescence a slender, lax, pendent, branched 
panicle of condensed cymes, up to 7 cm long, 
bearing 20-100 flowers; 3 flowers in clusters of 
5-10 along the axes, c. 2 by 2 mm; perianth lobes 
broad ovate-acute, c. 1-1 l / 2 by 1 mm, densely, short 
appressed-hairy outside; filaments l-172 mm > 
anthers ovoid, c. 1 mm 0; pistillode strongly 



rudimentary. — 5 Inflorescence a simple or 
branched panicle, up to 2'/ 2 cm long, 5-10- 
flowered; 9 flowers sessile along the axes, com- 
pressed ovoid-conical, 2-3 by IV2-2 mm; perianth 
lobes ovate-acute, densely appressed-hairy outside, 
17 2 -2 by 1 mm; ovary densely appressed-hairy, 
1V2-3 by 1-1 '/2 mm; stigmatic arms 5-10 mm. 
Infructescence up to 3 cm long, bearing (2-)4-6(-8) 
fruits. Fruit subglobose or ovoid, densely 
appressed-pubescent, 5-8 by 4-6 by 3-4 mm, 
short-beaked. 

Distr. Thailand; in Malesia: Malay Peninsula 
(incl. Singapore; common), Sumatra (rather rare), 
Borneo (common). 

Ecol. In primary and secondary forests, 0-1300 
m, mostly below 500 m ; often common locally as 
understorey tree in lowland forests. In Thailand it 
occurs mainly in the evergreen forest along streams. 
Fl. fr. Jan.-Dec., but mainly during July-Dec. 

Vern. Malaya: medang berbulu, m. kasap, tapis, 
M; N. Borneo: luazon, Kadasan, hugot-hugot, 
Dusun; Kalimantan: kayu ruas, Bandjar, gagas, 
Bassap. 

3. Gironniera hirta Ridl. J. Str. Br. R. As. Soc. n. 
82 (1920) 194; Fl. Mai. Pen. 3 (1924) 321; Soe- 
padmo in Whitmore, Tree Fl. Mai. 2 (1973) 417, 
f. 2. — Fig. 26o. 

Shrub to medium-sized tree up to 30 m, 30 cm 0. 
Bark smooth, light to grey-brown. Young branch- 
lets, petiole, stipules, terminal bud densely set with 
golden yellow, long, soft, erect hairs. Terminal 
buds ovoid-conical, 17 2 -3 by 74~7 2 cm; stipules 
F/2-3 by l j 2 cm. Leaves thick-coriaceous, rigid, 
ovate-elliptic or rarely elliptic-oblong, (6— )14— 18 
(-23) by (3— )5— 8(— 12) cm, index c. 2; base rounded, 
subcordate or attenuate, unequal-sided; margin 
entire, rarely distantly serrulate in the upper half; 
apex rounded, acute to acuminate; except for the 
midrib and nerves which are densely or sparsely 
set with long, appressed or erect, soft hairs, glab- 
rous above, densely soft-pubescent beneath; mid- 
rib and nerves raised and distinct beneath, flatfish 
or impressed above; nerves (8—) 1 0— 1 2(— 1 4) pairs, 
10-15 mm spaced, arcuating towards and anasto- 
mosing near the margin; reticulation subscalari- 
form to subareolate, well-spaced, distinct beneath, 
obscure above; petiole 2-10 by 1-3 mm, terete or 
flat above near the base. Inflorescences S or $, 
axillary, borne on separate branches, including the 
bracts densely golden yellow pubescent; bracts 
ovate-acute, 1-1 72 by 72-1 mm - — <S Inflorescence 
slender, lax, branched, thyrsoid, up to 8 cm long, 
bearing up to 100 flowers; <3 flower l l / 2 -2 mm 0, 
sessile along the axes, in cymoid clusters of 3-10; 
perianth lobes l'/ 2 -2 by 1 mm, densely appressed- 
hairy outside; filaments 1-1 72 mm » anthers ovoid- 
reniform, c. 1 mm ; pistillode strongly rudimen- 
tary. — ? Inflorescence (as seen in a very young 
infructescence) a branched panicle, up to 5 cm, 
carrying 2-20, sessile or short-stalked flowers; 
perianth lobes narrow ovate-acute, c. 2 by 1 mm, 
densely appressed short-hairy outside; ovary 
(young fruit) ovoid, densely short-, appressed- 
hairy, c. 3 by 2 mm; stigmatic arms up to 1 cm. 
Infructescence up to 10 cm long, axes c. 1 mm thick, 
bearing (2-)5-15(-20) fruits. Drupe ovoid-com- 
pressed, densely short-appressed-pubescent, 8-10 
by 6-8 by 3-5 mm, short-beaked. 



1977] 



Ulmaceae (Soepadmo) 



75 



Distr. Malesia: Malaya (rather rare), Sumatra 
(rare), Borneo (common), Moluccas (rare), New 
Guinea (rare, mainly in W.). 

Ecol. Scattered as an understorey shrub or tree 
in lowland forests, 0-700 m. Fl. fr. Jan.-Dec. 

Vern. Malaya & Sumatra: hampas tebu, 
hampilas burung, medang berbulu, m. kasap, M ; 
Kalimantan: kayu ruas, lempung bitlu, Bandjar; 
Sarawak: puloh, Iban; New Guinea: warpis, Biak. 

Note. Closely related to G. nervosa, but readily 
distinguished from the latter by the characters 
mentioned in the key. Specimens from Malaya 
(mainly from the Kluang area in Johore and 
Pahang), including the type, have much thinner 
leaves with a distantly serrulate margin than those 
from Sumatra, Borneo, and New Guinea. Since in 
most cases the field notes of the Malayan speci- 
mens indicate that the height of the tree was never 
more than 3 m, it is assumed that these specimens 
have been collected from saplings. 

4. Gironniera subaequalis Planch. Ann. Sc. Nat. 
Ill, 10 (1848) 339, p.p., excl. var. ceylanica; Bl. 
Mus. Bot. 2 (1856) 73, incl. var. brevistylis Bl., 
var. scabrida Bl. et var. serrulata Bl. I.e. 74; MlQ. 
Fl. Ind. Bat. 1, 2 (1859) 222; Planch, in DC. 
Prod. 17 (1873) 206; Hook./. Fl. Br. Ind. 5 (1888) 
485; Hemsl. J. Linn. Soc. Bot. 26 (1894) 452; 
Brandis, Ind. Trees (1906) 596; J. J. Smith in K. & 
V. Bijdr. 12 (1910) 666; Nova Guinea 8 (1912) 892, 
incl. var. papuana J. J. S. ; Laut. Bot. Jahrb. 50 
(1913) 326; Merr. En. Born. (1921) 217; En. 
Philip. 2 (1923) 35; Ridl. Fl. Mai. Pen. 3 (1924) 
320; Gagnep. Fl. Gen. I.-C. 5 (1927) 678; Corner, 
Ways. Trees (1940) 690; Back. & Bakh. /. Fl. 
Java 2 (1965) 12; Soepadmo in Whitmore, Tree Fl. 
Mai. 2 (1973) 419, f. 2. — Sponia annulata Teysm. 
& Binn. Ned. Kruidk. Arch. 3 (1855) 408. — G. 
costata Miq. in Zoll. Syst. Verz. (1855) 88; Fl. Ind. 
Bat. 1, 2 (1859) 223. — G. chinensis Bth. Fl. 
Hongk. (1861) 324. — G. nervosa var. subaequalis 
(Planch.) Kurz, For. Fl. Burma 2 (1877) 470. — 
G. amboinensis Laut. Bot. Jahrb. 50 (1913) 326. — 
G. longifolia Craib, Kew Bull. (1918) 371. — G. 
sumatrana Gandog. Bull. Soc. Bot. Fr. 66 (1919) 
288. — G. blumei Gandog. I.e. 288. — G. borneen- 
sis Gandog. I.e. 288. — G. ferruginea Gandog. 
I.e. 289. — Fig. 26h, m, p-r. 

Small to large-sized tree up to 40 m, 60 cm 0. 
Bark smooth to finely fissured, pustular or lenti- 
cellate. Terminal bud 2-3 by '/r'/icm; stipules 
linear-lanceolate, 1 V2-2V2 by l U- l l 2 cm. Leaves 
thick-coriaceous, broad ovate-elliptic or elliptic- 
oblong, (6-)12-16(-2l) by (3'/ 2 -)5-8(-13) cm, 
index 2-2' 2 , except for midrib and nerves glabrous; 
base attenuate or rounded, unequal-sided; margin 
entire or occasionally, especially when young, 
distantly serrulate at least for the upper half; apex 
rounded or attenuate-acute; midrib and nerves 
raised beneath, flattish above, sparsely or densely 
yellowish-brown appressed short-hairy beneath; 
nerves (6 )Kf 10) pairs. 10 I 5 mm spaced, at more 
than 60 with the midrib, arcuating and anasto- 
mosing along the margin; reticulations lubscalari- 
form, fine, dense or rather well-spaced, slightly 
and clearly visible beneath, flattish anil faintly 
visible above or obscure; petiole S I "> by I 2 mm, 
terete or adaxially Mat near the base, sparsely or 
densely appressed, simple, short, yellowish-brown 



pubescent, glabrescent. Inflorescence c? or ?, 
axillary, borne on separate branchlets or rarely on 
the same branchlet, including the bracts sparsely 
to densely short, yellowish-brown, appressed-hairy, 
glabrescent; bracts ovate-acute, 1-2 by 1 / 2 -l mm. 
— 3 Inflorescence paniculate, pendent, much- 
branched, 40-100-flowered, axes up to 3-7 cm 
long, V2-I mm thick; c? flowers l l / 2 -2 l j 2 mm 0, 
sessile and solitary along the axes or in clusters of 
3-5 on short, condensed secondary branches of the 
panicle; perianth lobes sparsely short appressed- 
pubescent, glabrescent, broadly ovate-rounded, 
2-2V 2 by l 1 / 2 -2 mm; filaments P/2-2 mm, anthers 
ovoid-subreniform, c. 1 by 1 mm; pistillode 
strongly rudimentary. — $ Inflorescence racemose, 
unbranched or more commonly branched, axes 
3-6 cm long, l-l'^rnm thick, bearing (2— )5— 10 
(-15) flowers; 9 flowers solitary and short-stalked 
along the axes, 2-A by 2 mm ; perianth lobes 
broadly ovate-acute, IV2-2 by l'^rnm, sparsely 
appressed-pubescent outside ; ovary 2-3 by 2 mm, 
densely appressed-pubescent, glabrescent; stig- 
matic arms up to 2 cm. Infructescence with a sturdy 
axis up to 2 mm thick, 5- 10 cm long, bearing (2-)5-8 
(-10) fruits. Fruit 1—1 */ 4 by 8-9 by 5-6 mm, sparsely 
appressed-pubescent, glabrescent, beak 2-5 mm. 

Distr. A rather variable species widely distri- 
buted in the Andaman Is., Burma, China (Yunnan, 
Kwangtung, Canton, Hainan), Hongkong, Indo- 
China, Thailand, throughout Malesia (except the 
Lesser Sunda Is.). 

Ecol. Understorey shrub or tree in primary and 
secondary forest, 0-1300 m, more commonly in 
the lowland between 200-500 m. Fl. fr. Jan.-Dec. 

Vern. Malaya: hampas tibu, m$dang kasap, M ; 
Sumatra: silu, siluk, M; W. Java: ki bulu, S; N. 
Borneo: kuayun, ruwayon, Dusun, untoh bulu, 
Iban; Kalimantan: katul, Bulungan; Anambas 
Is.: pupoh, M; W. New Guinea: gawa, giwa, 
migawa, Mooi, nadjun, nitjun, Kebar, bobohufeka , 
Manikiong, warpis, Biak. 

5. Gironniera parvifolia Planch. Ann. Sc. Nat. 
Ill, 10 (1848) 338 Cparvifolium')- Miq. Fl. Ind. 
Bat. 1, 2 (1859) 223; Planch, in DC. Prod. 17 
(1873) 206; Hook./ Fl. Br. Ind. 5 (1888) 486; 
Ridl. Fl. Mai. Pen. 3 (1924) 321 ; Corner, Ways. 
Trees (1940) 689; Soepadmo in Whitmore, Tree 
Fl. Mai. 2 (1973) 419, f. 2. — G. subaequalis var. 
ceylanica Planch. Ann. Sc. Nat. Ill, 10 (1848) 
339; Thw. En. PI. Zeyl. (1861) 268; Hook./ 
Fl. Br. Ind. 5 (1888) 485. — Helminthospeimum 
scabridum Thw. in Hook. J. Bot. Kew Misc. 6 
(1854) 303, t. 9c. — G. pauctnervta Merr. J. Str. 
Br. R. As. Soc. n. 77 (1917) 189; En. Born. 
(1921) 217. — G. zeylanica Gandog. Bull. Soc. 
Bot. Fr. 66 (1919) 288. — G. scabrida (Thw.) 
Alston in Trimen, Fl. Ceyl. 6 (1931) 267. — Fig. 
26k, n. 

Shrub M small-si/ed tree up to 15 m, 20cm 0. 

Bark smooth to finely fissured, lenticellate, grey- 
green or grey-brown, rerminal bud (5 )X I'm 18) 

by I 3 mm; stipules linear-acute, S 15 by 2 3 mm. 

Leaves chartaceous to thin-coriaceous, rarely 
coriaceous, elliptic-lanceolate <>i elliptic-obovate, 
irdy ovate-elliptic, (4 >* I2< 16) by (l*/i I 
1 k 5) cm, index * r 1, glabrous,; base attenuate 
mi rounded, more 01 less equal-tided; margin 

entire OI sometimes minutely and distantly sciiulale 



76 



Flora Malesiana 



[ser. I, vol. 8 2 



in the upper half (extra-Mai. and young speci- 
mens); apex attenuate-acute or rounded- 
acuminate; midrib and nerves slightly raised 
beneath, flattish above; nerves (5-)6-8(-9) pairs, 
7-10 mm spaced, at up to 60° with the midrib, 
arcuating and anastomosing along the margin; 
reticulations subareolate, fine, dense, visible 
beneath, obscure above; petiole 5-7 by 1-1 */ 2 mm, 
sulcate. Inflorescences $ or $, axillary, borne on 
separate branchlets, including the bracts sparsely, 
short, appressed-pubescent, glabrescent; bracts 
ovate-acute, membranous, c. 1 by l j 2 mm. — 
3 Inflorescence a slender, pendulous, simple or 
branched raceme, bearing 15-30 flowers, axes up 
to 5 cm long, 0.2-0.3 mm thick; <3 flowers solitary 
along the axes, short-stalked, l'/r^mm 0; 
perianth lobes broad ovate, c. l l l 2 -2 by l-l 1 /? mm; 
filaments 1-1 1 I 2 mm long, anthers ovoid-reniform, 
3 / 4 -l mm ; pistillode strongly rudimentary. — ? 
Inflorescence a slender, l-3(-5)-flowered raceme, 
4-10 cm long, axes 0.2-0.3 mm thick; $ flower 2-3 
by 2 mm, short-stalked; perianth lobes mostly 4, 
unequal in size, l l l 2 -2 l l 2 by 1-2 mm ; ovary sparsely 
short appressed-pubescent, 2-3 by 2 mm ; stig- 
matic arms up to IV2 mm. Infructescences slender, 
up to 10 cm long, bearing 1— 3(— 5) fruits, axes thin- 
ner than 1 mm. Fruit c. 8 by 6 by 4 mm, sparsely, 
short appressed-pubescent, glabrescent; beak up 
to 5 mm. 

Distr. Ceylon; in Malesia: Malay Peninsula 
(incl. Singapore; very common), Sumatra (rare), 
Borneo (common). 

Ecol. In primary as well as in secondary forest, 
0-1300 m, but mostly in the lowland, on various 
types of soil including those derived from lime- 
stone. Fl. Jan.-April, fr. Oct. -Dec. 

Vern. Malaya: hampas tebu, medang kasap, M; 
Sarawak: tepade, Kelabit. 

Note. As in other species of the genus, most of 
the specimens so far available suggest that the 
plant is monoecious. However, in Maingay K.D. 
1470 from Malaya both the $ inflorescence and 
infructescence are found on separate branchlets of 
the same collection number. Assuming that these 
branchlets were collected from the same tree, it 
would suggest that the species is monoecious but 
producing S and $ flowers at different stages of its 
growth. More field work is required to determine 
its breeding system. 

6. Gironniera rhamnifolia Bl. Mus. Bot. 2 (1856) 
74, t. 25; Miq. Fl. Ind. Bat. 1, 2 (1859) 223; 
Planch, in DC. Prod. 17 (1873) 206; Laut. Bot. 
Jahrb. 50 (1913) 326. — Fig. 261. 



Shrub to medium-sized tree, up to 25 m, 40 cm 
0. Bark smooth, pale grey-brown. Branchlets 
initially densely set with simple, yellowish-brown, 
soft, erect hairs, glabrescent. Terminal buds 1-1 V2 
by 0.2-0.3 cm ; stipules narrow elliptic-lanceolate, 
c. IV2 by l U cm. Leaves chartaceous to thin- 
coriaceous, glabrous, elliptic-lanceolate or rarely 
ovate-elliptic, (5-)8-15(-17) by (2-)4-6(-8) cm, 
index 2 l l 2 -VI 2 ; base rounded or attenuate, ± 
equal-sided; margin entire; apex acute to acumi- 
nate; midrib and nerves slightly raised beneath, 
flattish to slightly impressed above, often densely, 
short, appressed-hairy; nerves 10-13 pairs, at an 
angle of over 60° with the midrib, arcuating and 
anastomosing towards the margin; reticulations 
fine, lax, subscalariform to subareolate, faintly 
visible beneath; petiole 5-10 by l-l'^mm, 
glabrous, shallowly furrowed or flat above near the 
base. Inflorescences <3 or ?, axillary, including the 
bracts sparsely short, yellowish-brown appressed- 
pubescent; bracts narrow ovate-acute, c. 1 by 
V2 mm. — c? Inflorescence an unbranched or 
branched panicle, 3-5 cm long, bearing 15-30 
flowers; <$ flowers sessile and solitary along the 
axes, c. 1-1 '/2 mm 0; perianth lobes ovate, 
IV2-2 by 1 mm, sparsely short, appressed- 
pubescent outside; filaments l l / 2 -2mm, anthers 
ovoid-reniform, l j 2 -\ mm 0; pistillode strongly 
rudimentary. — $ Inflorescence an unbranched, 
(l-)3-5-flowered panicle, 3-4 cm long with the 
axes c. V2-I mm thick; ? flowers 2-3 by 2 mm; 
perianth lobes 5, ovate, IV2-2 by 1 mm; ovary 2-3 
by 2 mm, densely short, appressed-pubescent, 
glabrescent; stigmatic arms up to l'^cm. Infruc- 
tescence up to 5 cm long, axes c. 1 mm or thicker, 
bearing (l-)3-5 fruits. Fruit ± glabrous, c. 7 by 6 
by 3 mm, short-beaked. 

Distr. Malesia: N. Moluccas (Morotai), New 
Guinea (incl. Biak, Japen, and Polima Is.). 

Ecol. Primary and secondary forests, locally 
often very common in rocky or stony habitats 
including limestone, 0-1000 m. Fl. fr. Jan.-Dec. 

Vern. W. New Guinea: warpis, Biak, tamanpara, 
Japen, rami, Iko. 



Excluded 

Gironniera glabra Merr. Philip. J. Sc. 1 (1906) 
Suppl. 42 = Paratrophis glabra Steen. J. Bot. 72 
(1934) 8 = Chevalierodendron glabrum Leroy, 
C. R. Ac. Sc. Paris 227 (1948) 146 = Streblus 
glaber (Merr.) Corner, Gard. Bull. Sing. 19 
(1962) 221 (Moraceae). 



IRIDACEAE (D. J. L. Geerinck, Bruxelles) 1 

Perennial herbs, often with bulbs, tubers or rhizomes, sometimes undershrubs. 
Leaves simple, equitant (except in Crocus), with parallel nerves. Inflorescences 
terminal or axillary, in cymes, spikes or panicles, sometimes very contracted or 
flowers solitary, bracteate and with 1 or 2 spathes. Flowers bisexual, actinomorphic 
to zygomorphic, often marcescent. Tepals free or united into a tube, in 2 whorls, 
the inner ones rarely inconspicuous (Patersonia). Stamens 3 or exceptionally 2 with 
1 staminode (in the Australian Diplarrhena), free or united into a tube, basifixed 
or dorsifixed, opposite to the outer tepals. Ovary inferior (or superior in the 
Tasmanian Isophysis), 3-celled with axillary placentas; style entire or trifid, 
sometimes tepaloid; stigmas 3 or 6, terminal or sometimes axillary, alternating 
with or opposite to the outer tepals; ovules generally numerous. Fruit capsular, 
dehiscing loculicidally, apically or irregularly. Seeds angular, flat or globose, some- 
times winged. 

Distribution. Cosmopolitan, with c. 60 genera and c. 800 spp., predominantly in the tropics and the 
southern hemisphere. In Malesia: only two Australasian genera each with 1 sp., and four exotic ones 
introduced and naturalized. 

Many are cultivated in botanic gardens and occasionally in private gardens; see for an elaborate 
treatment Backer, Handb. Fl. Java 3 (1924) 116-130 and Backer & Bakh./. Fl. Java 3 (1968) 144-154. 

Ecology. Both native species are characteristic mountain plants. 

Morphology. This family is usually herbaceous, but in a few genera (Klattia, Nivenia, Patersonia, 
Witsenia) stems may be woody at the base. The leaves are equitant and are laterally compressed, the two 
halves are free at the sheathing base and gradually fused to the top. The flowers are actinomorphic to 
distinctly zygomorphic with intermediate forms, sometimes in the same genus. 

Uses. Belamcanda chinensis and Eleutherine palmifolia are used for medicinal purposes, probably mainly 
on account of their glucosides; cf. Heyne, Nutt. PI. (1927); Burkill, Diet. (1935); Quisumbing, Med. PI. 
Philip. (1951). 

KEY TO THE GENERA 

1. Flowers all sessile. Tepals united into a tube. Capsules included in bracts or spathes. 
2. Flowers actinomorphic. Inner tepals inconspicuous. Stamens united into an undivided or trifid tube. 
Caespitose or rhizomatous herbs or undershrubs 1. Patersonia 

2. Flowers ^ zygomorphic. Tepals unequal. Stamens free. Cormogenous herbs. . . . 5. Gladiolus 
1. Flowers all pedicelled. Tepals free or nearly so. Capsules exserted from bracts or spathes. 

3. Tepals clawed. Stamens free. 

4. Cormogenous herbs. Stamens appressed against the back of the style-arms. Ovary not beaked. 
Style trifid with bilobed arms 6. Trimezia 

4. Rhizomatous herbs. Stamens not appressed against the back of the style-arms. Ovary beaked. 
Style trifid with undivided arms 3. Belamcanda 

3. Tepals not clawed. 

5. Tepals shortly connate, subequal. Stamens united into a tube, rarely nearly free. Caespitose to 
rhizomatous herbs 2. Sisyrinchium 

5. Tepals free, the inner ones smaller. Stamens free. Bulbous herbs 4. Eleutherine 

1. PATERSONIA 

R.Br, ex Kfr-Gawl. Bot. Mag. (1807) t. 1041, nom. cons.; Prod. Nov. Holl. 
(1810) 304; Bth. Fl. Austr. 6 (1875) 400; Ghfrinck, Bull. Jard. Bot. Nat. Belg. 44 
(1974) 41. - Genosiris Lakh i.. Nov. Holl. PI. Sp. 1 (1804) 13, t. 9. — Fig. 1-3. 

Caespitose to rhi/omatous herbs or undershrubs. Inflorescences terminal, in 
few-flowered contracted cymes, each with 2 spathes. /'lowers actinomorphic, 
sessile, bluish to purple, exceptionally yellow or whitish, Tepals dimorphic, united 
into a long and filiform tube at the base, the inner lobes inconspicuous. Stamens 3, 

(l) With co-operation by the General I dhor. 

(77) 



78 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 1. Patersonia lowii Stapf on Mt Losir, Gajolands, N. Sumatra, at c. 2400 m altitude (Photogr. de 

Wilde-Duyfjes, April 1975, n. 16390). 



1977] 



Iridaceae (Geerinck) 



79 



united into an undivided or trifid tube. Ovary cylindrical, lanate; style entire; 
stigmas 3, subfoliaceous, alternating with the outer tepals. Capsules loculicidal, 
included. Seeds angular or ellipsoidal. 

Distr. Australia and Tasmania (12 spp.), and Malesia (1 sp.). 

Ecol. Open, low shrubberies, heaths and sedge-lands, 2000-3500 m. 

Note. In sterile state the habit of Patersonia is strikingly resembling that of the sedge genus Machaerina. 
A specimen mentioned by Went/. (Nova Guinea 14, 1924, 1 14) as Patersonia from Mt Goliath (de Kock 
50) belongs to Machaerina, as corroborated anatomically by Dr P. Baas. 



I. Patersonia lowii Stapf, Trans. Linn. Soc. Bot. 

II, 4 (1894) 241, t. 20, f. 7-9; Merr. Philip. J. Sc. 2 
(1907) Bot. 268; En. Born. (1921) 119; En. Philip. 
1 (1923) 220; Not. Nat. Ac. Nat. Sc. Philad. n. 47 
(1940) 2. — P. borneensis Stapf, Trans. Linn. Soc. 
Bot. II, 4 (1894) 241 ; Gibbs, J. Linn. Soc. Bot. 42 
(1914) 165; Merr. En. Born. (1921) 1 \9. — P. novo- 
guineensis Gibbs, Arfak (1917) 101 ; Went/. Nova 
Guinea 14 (1924) 114, incl. var. auriculata Went, 
I.e.; Hatus. Bot. Mag. Tokyo 56 (1942) 426. — 
Fig. 1-3. 

Tufted herb, 15-60 cm high. Leaves basal to 
subbasal, fiat to — biconvex, 5-60 cm by 3-6 mm, 




reddish or rarely whitish tomentellous to glab- 
rescent along the margins towards the top, ± 
glaucous. Inflorescences equalling the leaves or 
nearly so; peduncle 8-50 cm long, glabrous, the 
lower part surrounded by a persistent central leaf; 
spathes suboval to narrowly suboval, 2*/2-5 cm by 
7-12 mm, dark brown-orange, greyish when 
growing old, distinctly striate, with a red-hairy line 
on the keel to glabrous. Flowers bluish to pale 
mauve or purple, sometimes whitish; perigone- 
tube 2-2'/2 cm long, the outer lobes 8-16 by 6-10 
mm. Staminal tube entire; anthers yellow. Ovary 
c. 5 mm long. Capsules 2-3 cm long ; valves 3-4 mm 
wide. Seeds c. 2 mm, black. 

Distr. Malesia: Sumatra (Gajolands: Mt Losir), 
Borneo (Mts Kinabalu and Murud, Kalabit 
Highlands), Philippines (Mindoro), New Guinea 
(Tamrau Range, Arfak Mts, Mamberamo River, 
Central to Milne Bay Districts). Fig. 4. 

Ecol. Open shrubby vegetation or open forests, 
sedge meadows and heaths, on stony or impervious 



Fig. 2. Patersonia lowii Stapk. a Habit, b. capsule, 

both nat. size, c. seed, • 5 (a van Royen &. 

Si i umer 7102. b c Brass 22259). 




Fig. 3. Patersonia lowii Si aim. Same locality as in 
fig. 1. 



80 



Flora Malesiana 



[ser. I, vol. 8 2 



clay soils, often gregarious, 2000-3500 m. Fl. fr. 
Dec.-Aug. Flowers open early in the morning but 
become soon marcescent. 

Vern. New Guinea: atetdzjii, Mt Arfak, 
Manikiang lang. 

Notes. The sizes of the tepals and of the seeds 
have been taken from the original descriptions of 
the synonymous taxa. 



A variable species concerning the indument; the 
disjunct populations are not uniform. It seems to 
be allied to the widely distributed Australian 
P. fragilis (Labill.) Ascherson & Graebner, 
which differs by glabrous leaves and spathes, 
inflorescences much shorter than the leaves and the 
lower part of the peduncles never surrounded by a 
central leaf. 




^-i 







Fig. 4. Range of Pater sonia lowii Stapf. 



2. SISYRINCHIUM 



Linne, Sp. PL (1753) 954; Johnston, J. Am. Arb. 19 (1938) 376; Foster, Contr. 
Gray Herb. 166 (1948) 28. — Renealmia R.Br. Prod. (1810) 592, pro parte, non L. 
/. 1781. —Fig. 5-6. 

Caespitose to rhizomatous herbs. Inflorescences axillary or terminal in panicles 
of fan-shaped and few-flowered cymes or of many-flowered clusters, each with 1-2 
spathes. Flowers actinomorphic, pedicelled, bluish or yellowish. Tepals subequal, 
shortly connate at the base. Stamens 3, united into a tube at the base, rarely nearly 
free. Ovary: style trifid, the lobes filiform; stigmas 3, small, alternating with the 
outer tepals. Capsules exserted. Seeds small, globose. 



Distr. About 100 spp. in Central and South America, 1 native sp. in New Zealand, Australia and East 
Malesia (New Guinea); also one species introduced. 

The Papuan-Australasian species was mostly arranged in the closely allied genus Libertia Spr. How- 
ever, the tepals are not dimorphic but about similar so that it must be arranged in Sisyrinchium. 



1977] 



Iridaceae (Geerinck) 



81 



KEY TO THE SPECIES 



Leaves cauline, distichous. Inflorescences in 1-2-flowered, contracted cymes, a few at a cauline leaf. 

Flowers c. 5 mm long. Stamens nearly free, glabrous 1. S. pulchellum 

Leaves mostly basal. Inflorescences in 3-6-flowered cymes, generally 2 at a cauline leaf. Flowers c. 7 mm 
long. Stamens united in a hairy tube, trifid at the top 2. S. micranthum 



1. Sisvrinchium pulchellum (R.Br.) F.v.M. Fragm. 
Phyt. Austr. 7 (1870) 92; Trans. R. Soc. Victoria 1 
(1889) 34; Geerinck, Bull. Jard. Bot. Nat. Belg. 44 
(1974) 59. — Renealmia pulchella R.Br. Prod 
(1810) 592, to replace S. pulchellum R.Br. I.e. 305 
— Libertia pulchella Spr. Syst. Veg. 1 (1824) 169 
Lane-Poole, For. Res. (1925) 77; Laut. Bot 
Jahrb. 62 (1929) 462; Steen. Bull. Jard. Bot 
Btzg III, 13 (1934) 220; Hoogl. Blumea 4 (1958) 
235; Balgooy, Pac. PI. Areas 2 (1966) 286; 
L. Moore, New Zeal. J. Bot. 5 (1967) 267. — Fig. 
5a. 

Glabrous herb, 10-35 cm high. Leaves cauline, 
distichous, linear, 4-22 cm by 1-10 mm. Inflores- 
cences in 1-2-flowered, contracted cymes, a few at a 
cauline leaf, each cyme with 1 spathe: this 4-15 
(-40) by 2-6 mm. Pedicels to 4 cm. Flowers c. 
5 mm long. Tepals white to yellowish, c. 4 by 
l 1 2 -2 mm. Stamens nearly free, filaments c. 4 mm, 
anthers 1 mm long. Ovary ellipsoid, c. 1 mm long; 
style with undivided part 1-2 mm, the lobes 1-2 
mm long. Capsules globular, 2-5 mm ; valves 
c. 2 mm wide. Seeds black, 1 mm 0. 

Distr. New Zealand, Australia (New South 
Wales, Victoria, Tasmania), and East Malesia: 
New Guinea (Lake Habbema area and Mt Antares 
in West, many localities in East). Fig. 6. 




Ecol. Open forests and shrubby vegetation, in 
tree fern heath and alpine grassland, common on 
Mt Sarawaket in Libocedrus-Dacrydium forest 
(Lane-Poole), 2400-3700 m. Fl. mostly Jan.-Aug. 

Note. Moore (I.e. 255-275) studied the variation 
in New Zealand and distinguished three species 
among which are two polyploids. 




Fig. 5. Siwriw hium pulchellum (R.Hk.) F.V.M. a. 
Inflorescence, nat. size. S micranthum Cav. 

b. Inflorescence, nat. m/c (a Kai kman 4497, b van 
Royen 16028). 



Fig. 6. Range of Sisyrinchium pulchellum (R.Br.) 
F.v.M. 



2. Sisyrinchium micranthum Cav. Diss. Bot. 6 
(1788) 345, t. 191 ; Back. Handb. Fl. Java 3 (1924) 
125; Johnston, J. Am. Arb. 19 (1938) 390; 
Foster, Contr. Gray Herb. 166 (1948) 31 ; Willis, 
Handb. Fl. Victoria 1 (1962) 335 ; Steen. Blumea 15 
(1967) 154; Back. & Bakh. /. Fl. Java 3 (1968) 
150. — Fig. 5b. 

Slender herb, 5-25 cm high, glabrous, with a 
flat stem. Leaves few, mostly basal, linear, 3-12 cm 
by 1-5 mm. Inflorescences in 3-6-flowered cymes, 
generally 2 at a cauline leaf, each cyme with 2 
spathes, outer spathe 20-30 mm long, inner 
spathe 15-25 mm long, both 1-2 mm wide. Flowers 
c. 7 mm long. Tepals yellow with red or brown 
markings, c. 6 by 1 mm. Stamens united in a hairy 
tube, trifid at the top; filaments c. 1 mm long; 
anthers l /a mm ' or >g- Ovary ellipsoid, laxly hairy, 
c. 1 mm long; style with the undivided part c. ' mm 
long, the lobes c. l l 2 - 3 U mm long. Capsules 
globose, 2-3 mm long; valves c. 2 mm wide. 
Seeds black, 1 mm 0. 

Distr. Southern to Central America ; naturalized 
in Australia, New Caledonia, Fiji, New Zealand, 
and also in Malesia: W. Java (Tjibodas), E. New 
Guinea (Morobe Distr.: Edie Creek), perhaps 
elsewhere. 

Ecol. In the vicinity of the Tjibodas Botanic 
Garden as a weed in grassland and waste places, at 
c. 1450 m, certainly escaped from the garden. In 
Papua at c. 1950m almost certainly introduced 
from Australia where it was first recorded about 
1870. Poisonous to stock. 



82 



Flora Malesiana 



[ser. I, vol. 8 2 



3. BELAMCANDA 

Adans. Fam. 2 (1763) 60. 

Rhizomatous herbs. Inflorescences terminal, in panicles of flowered and corym- 
biform cymes, each with 2 spathes. Flowers actinomorphic, pedicelled, yellowish to 
orange. Tepals subequal, shortly connate, clawed. Stamens free. Ovary beaked; 
style trifid, the lobes short; stigmas 3, small, alternating with the outer tepals. 
Capsules exserted. Seeds large, globose. 

Distr. Monotypic, native of China and Japan, cultivated in Malesia and many other countries and 
sometimes naturalized. 



1. Belamcanda chinensis (L.) DC. in Redoute, 
Liliac. 3 (1805) t. 121 ; Koord. Minah. (1898) 313; 
C. B. Rob. Philip. J. Sc. 6 (1911) Bot. 196; Koord. 
Exk. Fl. Java 1 (1911) 312; Merr. Fl. Manila 
(1912) 152; Philip. J. Sc. 11 (1916) Bot. 260; En. 
Philip. 1 (1923) 220; Back. Handb. Fl. Java 3 
(1924) 124; Heyne, Nutt, PI. (1927) 461 ; Gagnep. 
Fl. Gen. I.-C. 6(1934) 675; Burk. Diet. (1935) 315; 
Steen. Fl. Sch. Indon. (1949) 147; Quis. Med. 
PI. Philip. (1951) 181; Henders. Mai. Wild Fl. 
Monoc. (1954) 168, f. 98; Ohwi, Fl. Japan (1965) 
316; Back. & Bakh. /. Fl. Java 3 (1968) 149. — 
Balem-canda schularmani Rheede, Hort. Mai. 1 1 : 
73, t. 37. — Ixia chinensis Linne, Sp. PI. (1753) 
36. — Pardanthus chinensis Ker-Gawl. in Konig 
& Sims, Ann. Bot. 1 (1805) 247; Bl. En. PI. Jav. 
(1827) 26; Zoll. Syst. Verz. 1 (1854) 70; Filet, 
PI. Bot. Tuin Weltevreden (1855) 13; Miq. Fl. Ind. 
Bat. 3 (1859) 579; Blanco, Fl. Filip. ed. 3 (1877- 
83) t. 376. 

Corymbosely branched, glabrous herb, 1-1 Vi m 
high. Leaves distichous, basal and cauline, broadly 
linear, 30-60 by 2-4 cm, glaucous. Inflorescences 
in 6-12-flowered cymes, once or twice branched; 
spathes membranous, c. 10 by 4 mm. Pedicels 
2-4 cm, persistent. Tepals yellowish to orange, 
with red spots, unguiculate, obovate, 25-35 by 



c. 7 mm, outer ones largest. Filaments filiform; 
anthers c. 10 mm long. Ovary ellipsoid, c. 5 mm 
long; style-arms gradually thickened upwards. 
Capsules obovate, 15-20 mm long; valves 8-12 mm 
wide. Seeds shining black, 5 mm 0. 

Distr. Native in China and Japan; in Malesia 
introduced and cultivated, locally naturalized 
(Sumatra, Java, S. Celebes, Philippines, Moluccas: 
Morotai, Banda, Ceram); cultivated and locally 
naturalized in many tropical and subtropical coun- 
tries, e.g. Hainan, Taiwan (Formosa), Tonga, etc. 

Ecol. In Java it is mainly naturalized in the 
eastern part between 750 and 2100 m, occurring in 
thickets and forest edges, and said not to grow 
well at low altitude. 

The flowers open in the forenoon and have 
withered by midday. 

Uses. Heyne and Burkill (ll.ee.) mention usage 
for several minor medicinal purposes, the dried 
rhizome being used as a purgative and for com- 
plaints of the chest and liver, etc. 

Vern. Sumatra: piso-piso, Batak. Java: akar 
tjamaka, djamaka, gegebangan brodjo lintang, 
suliga, S, semprit, wordi, J. Celebes: karimenga 
kulo, katna, ketep, ketew, kiris, Minahasa, Alf. 
lang., tagari, Bonthain. Philippines: abinaco. 



4. ELEUTHERINE 

Herbert, Bot. Reg. 29 (1843) t. 57, nom. cons. 

Bulbous herbs. Inflorescences axillary in few-flowered and contracted cymes, 
each in 2 spathes. Flowers actinomorphic, pedicelled, whitish. Tepals free, the inner 
ones smaller. Stamens 3, free. Style deeply trifid; stigmas 3, small, alternating with 
the outer tepals. Capsule loculicidal, exsert. Seeds ellipsoidal to angular. 

Distr. In America 2 spp. and according to Gagnepain (Fl. Gen. I.-C. 6, 1934, 676) 2 spp. in Indo- 
China. Introduced in Malesia, and locally naturalized. 



1. Eleutherine palmifolia (L.) Merr. Philip. J. Sc. 7 
(1912) Bot. 233; Fl. Manila (1912) 153; Sp. Blanc. 
(1918) 104; En. Philip. 1 (1923) 220; Quis. Med. 
PI. Philip. (1951) 182; Back. & Bakh./. Fl. Java 3 
(1968) 150. — Sisyrinchium palmifolium Linne, 
Mant. 1 (1767) 122. — Sisyrinchium bulbosum 
Mill. Gard. Diet. ed. 8 (1768) n. 3. — Ixia 
americana Aubl. PI. Guian. 1 (1775) 33. — Moraea 
plicata Sw. Fl. Ind. Occ. 1 (1797) 82. — Antholyza 
meriana (non L.) Blanco, Fl. Filip. (1837) 24; ed. 3, 



1 (1877) t. 100; Merr. Publ. Gov. Lab. Philip. 27 
(1905) 85. — E. plicata Herbert, Bot. Reg. 29 
(1843) t. 57. — E. bulbosa (Mill.) Urban in 
Fedde, Rep. 15 (1918) 305; Leonard, Bull. Soc. 
R. Bot. Belg. 84 (1951) 55. — E. americana Merr. 
ex Heyne, Nutt. PI. Ned. Ind. ed. 2, 1 (1922) 502; 
Back. & Sloot. Handb. Thee (1924) 91, t. 91; 
Back. Handb. Fl. Java 3 (1924) 126; Heyne, Nutt. 
PI. (1927) 462. 
Herb, 30-60 cm high, glabrous; bulb red, ovoid, 



1977] 



Iridaceae (Geerinck) 



83 



2V2-5 cm long. Leaves basal 3^4 from each bulb, 
and cauline, narrowly elliptic, plicate-nerved, 
25-60 by 1-2 1 2 cm. Inflorescences in 4-10-flowered 
cymes; spathes 12-16 mm long, green. Flowers 
very fugacious, white. Tepals obovate, c. 1 5 mm 
long. Stamens yellow to orange, 8-10 mm long. 
Ovary ellipsoidal, c. 2 mm long; style-arms 
filiform, yellow; stigmas white. Capsules globose, 
c. 6 mm long. Seeds dark brown, c. 2 mm 0. 

Distr. Native in tropical America, cultivated 
and naturalized in tropical Africa and in Malesia: 
W. Java, W. Borneo, and the Philippines (Luzon, 
Leyte, Negros, Mindanao); in Java already noticed 
± 1820. 

Ecol. A weed, finally tufted, in estates and waste 
places, which multiplied by its tubers; c. 150-1500 
m. In Java the scentless flowers open at about 5 
o'clock in the afternoon, but have already wilted at 
about 7 o'clock. 



Uses. According to Quisumbing I.e. in the 
Philippines macerated bulbs are applied on the 
stomachs of children to relieve gas pains, and a 
decoction is diuretic. According to Heyne I.e. this 
finally strongly stooling, tufted plant is cultivated 
and its bulbs have various applications in native 
medicine: diuretic, purgative, emetic, against 
dysentery, jaundice, etc. 

Vern. Vijfuursbloem, D. Java: babawangan, b. 
beureum, bawang sabrang, b. sieum, S, bawang 
kapal, M, brambang sabrang, luluwan sapi, tiki 
sabrang, J. Philippines: ahos-dhos, C. Bis., bakong 
sa Persia, mala-bauang, rosas sa Siam, Tag., hagu- 
sahis, S. L. Bis., palmilla, Spanish. 

Note. The tepals are sometimes numerous, up to 
15; the number of the stamens is sometimes up to 
8; the ovary is sometimes 4-11-locular with the 
same number of style-arms. 



5. GLADIOLUS 

Linne, Sp. PI. (1753) 36; Geerinck, Bull. Jard. Bot. Nat. Belg. 42 (1972) 269; 
Lewis & Obermeyer, J. S. Afr. Bot. Suppl. 10 (1972). 

Cormogenous herbs. Inflorescences terminal or sometimes axillary in spikes, 
rarely secund. Flowers ± zygomorphic, sessile, various in colour, each in a single 
spathe. Tepals unequal, united into an often curved tube. Stamens 3, often arched, 
free. Style entire; stigmas 3, obovate, alternating with the outer tepals. Capsule 
loculicidal, included. Seeds often winged. 

Distr. About 180 spp. in Africa, South Europe and West Asia, one locally naturalized in Malesia. 



1. Gladiolus natalensis (Ecklon) Reinw. ex Hook. 
Bot. Mag. (1831) t. 3084; Geerinck, Bull. Jard. 
Bot. Nat. Belg. 42 (1972) 281 ; Lewis & Obermeyer, 
J. S. Afr. Bot. Suppl. 10 (1972) 44. 

var. natalensis. 

Stems 50- 150 cm. Leaves almost basal, broadly 
linear, up to 30 by 4-5 cm. Inflorescences terminal 
in 2-25-flowered spikes; spathes 4-8 cm long. 
Flowers yellowish to pinkish, often with brown 



markings. Perigone-tube curved, 2-5 cm long, 
lobes unequal, the upper 4'/2-5 cm, the outer 
laterals 3-4 1 ! 2 cm, the inner laterals 2-3 cm and the 
lower 2 l l 2 -3 l l 2 cm long. Filaments curved, 4 l l 2 -6 
cm; anthers lV4-l 3 /4 cm. Ovary ellipsoid, 5-8 mm 
long; style curved, 2-7 cm; stigmas 5-7 mm. 
Capsule ellipsoid, 2-5 cm long; valves 7-10 mm 
wide. Seeds flat, winged, golden brown, 7-lOmm0. 
Distr. Tropical and southern Africa, naturalized 
in Malesia: Philippines (Luzon). 



6. TRIMEZIA 

Salisb. ex Herbert, Bot. Reg. 30, Misc. (1844) 88; Diels, Pfl. Fam. ed. 2, 15a 
(1930) 497; Foster, Rhodora 64 (1962) 307. 

Cormogenous herbs. Inflorescences axillary in few-flowered cymes, each in 2 
spathes. /'lowers actinomorphic, pedicelled, yellowish to reddish. Tepals free, 
clawed, the inner ones shorter and narrower with recurved tops. Stamens free, 
against the back of the style-arms. Style trifid, the lobes broadly flat, bilobed; 
stigmas small, opposite to the outer tepals. Capsules apieally dehiscent, with 3 pores 
exserted beyond the spathe. Seeds globose to angular. 



Distr. A few species m ( cntral and tropical America, one introduced in Malesia (Malaya, West Java) 



84 



Flora Malesiana 



[ser. I, vol. 8 2 



1. Trimezia martinicensis (Jacq.) Herbert, Bot. 
Reg. 30, Misc. (1844) 88; Back. Handb. Fl. Java 3 
(1924) 121 CTrimeza , )\ Henders. Mai. Wild Fl. 
Monoc. (1954) 168, f. 98; Foster, Rhodora 64 
(1962) 308; Back. & Bakh. /. Fl. Java 3 (1968) 
148. — Iris martinicensis Jacq. En. PI. Carib. 
(1760) 12. — T. lurida Salisb. Trans. Hort. Soc. 1 
(1812) 308; Henders. Gard. Bull. S. S. 4 (1928) 
341. — Cipura martinicensis Kth in H. B. K. Nov. 
Gen. Sp. 1 (1816) 320. 

Glabrous herb, 100-150 cm. Leaves basal to 
cauline, linear, 20-100 cm long and 8-12 mm wide. 
Inflorescences in 3-6-flowered cymes, solitary or 
binate; spathes 2-2'/ 2 cm long, 10—25 cm 
peduncled. Pedicels \ l l 2 -yi 2 cm- Flowers yellow, 
brownish at the base. Outer tepals obovate, erect 



to patent, 19-25 mm long and 10-13 mm wide; 
inner tepals narrower, S-shape curved. Stamens 
3-4 mm long. Ovary ellipsoidal ; style-arms 5-7 mm 
long, shortly bilobed. Capsule ellipsoid, 13-20 mm 
long. Seeds brown, superficially ribbed. 

Distr. Native of Mexico, cultivated and locally 
naturalized in Malesia: Malaya, West Java. 

Ecol. In sunny or slightly shaded localities, 
between grass, originally in Malaya at Kuala 
Lumpur, but now not uncommon in Malaya 
(Henderson), in West Java at Bogor on and around 
a native cemetery (Backer), in both cases escaped 
from a Botanic Garden, below 250 m. Flowers 
expand in the forenoon and have withered by 
midday. 

Vern. Forenoon yellow flag, E. 



CORNACEAE (K. M. Matthew, Tiruchirapalli, India) 1 

In the past century Cornaceae were mostly delimited in a wide sense and they 
represented a fairly heterogeneous assemblage. Harms (Ber. Deut. Bot. Ges. 15, 
1897, 28 and in E. & P. Nat. Pfl. Fam. 3, 8, 1898, 255) distinguished 7 subfamilies. 
Of these Garryoideae were later mostly recognized as a separate family Garryaceae, 
Alangioideae as Alangiaceae, Nyssoideae and Davidioideae together as Nyssaceae, 
leaving Cornaceae with the remaining three subfamilies Cornoideae, Curtisioideae 
(monotypic, South Africa) and Masti.xioideae (monotypic, Indo-Malesian tropics). 
Cf. Wangerin, Pfl. Reich Heft 41 4 (1910) 18. 

In recent years, however, the other genera (6) of the Cornoideae, besides Cornus, 
have also been recognized as monotypic families, with the exception of Corokia 
which was transferred to Saxifragaceae-Escallonioideae . Notably Takhtajan 
(Proiskh. Prokruitosem. Rast. : 89, non vidi) is in favour of these monotypic families. 
In his 'Flowering Plants' (ed. C. Jeffrey; 1969: 227) he accepted 7 segregate 
families besides Cornaceae sens. str. (omitting mention of two Madagascan genera, 
one of which he had formerly also raised to family rank, according to Shaw, 
1973). These 7 families he arranged, together with Araliaceae and Umbelliferae, in 
the order Cornales, a phylogenetic construction of affinity not much different from 
earlier conceptions. The general impression is thus that the distinction of the 
segregate families is largely an inflation in rank. 

We have not followed this tendency towards inflation advocated by a few con- 
temporary systematists and have accepted Cornaceae in the wide sense. We do not 
feel that inflation has the merit of improving scientific insight in the mutual syste- 
matical affinities, which remain as they were, either as tribes or as subfamilies, 
representing together one phylogenetical whole. In addition the disadvantage of the 
inflation is that the multiplication of family names becomes unnecessarily a real 
challenge to our capacity to memorize, and deflates firmly established family 
concepts. 

We briefly mention that further relationships are sometimes suggested with 
quite remote groups. Rendle (Class. Fl. PI. 2, 1952, 422) suggested alliance with 
Caprifoliaceae, e.g. Viburnum; affinity has also been suggested with Saxifragaceae- 
Escallonioideae. It falls beyond the scope of the present account to elaborate further 
the extensive literature on the subject. 

Cornaceae are in great majority northern extratropical, in which zone also many fossils are known. 
There are some stray genera on the southern hemisphere. Mastixia is tropical but was found in abundance 
in the Tertiary in the subtropics and warm-temperate regions of the northern hemisphere. See under the 
genus. 

Note. Besides the native genus Mastixia the family is represented in Java by Aucuba japonica Thunb. 
which is sometimes cultivated in the mountains. Cf. Back. & Bakh./. Fl. Java 3 (1965) 159. — Ed. 

1. MASTIXIA 

Blume, Bijdr. (1826) 654; Harms in E. & P. Nat. Pfl. Fam. 3, 8 (1898) 262; 
Wangerin, Pfl. Reich Heft 41 4 (1910) 19; Hall./. Beih. Bot. Ccntralbl. 34, 2 
(1916)40;Dansik, Blumea 1 (1934)47; Matthlw, Blumea 23 (1976) 51, f. 1-6. 
Fig. 1, 3. 

(I) ( < imposed from the precursory revision in Blumea 21 (1976) SI 93 by the General I ilitor. 

Thanks arc due l<>r the financial assistance by the Netherlands Organization for the Advancement of 
Pure Research (Z.W.O.). 

(85) 



86 



Flora Malesiana 



[ser. I, vol. 8 2 



Unarmed, resinous, evergreen trees up to 40(-60) m ; branchlets with pith. Leaves 
simple, exstipulate, entire, acute, alternate or (sub)opposite to decussate, sometimes 
with domatia. Thyrses terminal on the main shoots, sometimes also on the laterals, 
up to 4(-8) times branched, the branches of the first order either (sub)opposite 
(' Oppositae') or spirally arranged ('Alternae''); further branchings with a tendency 
towards decussate arrangement and terminated by cymes; cymes with the central 
flower most often sessile and ebracteolate, lateral flowers pedicelled and bracteo- 
late. Bracts and bracteoles ovate to triangular, connate or free, lower bracts 
sometimes gradually becoming foliaceous. Flowers bisexual, greenish to yellowish. 
Calyx 4-5(-6-7)-toothed or -lobed, persistent. Petals valvate, 4-5(-6), thick, ovate 
to oblong-elliptic, inflexed at apex and 2-dentate or fimbriate, sometimes with a 




Fig. 1. Mastixia kaniensis Melch. ssp. kaniensis. a. Habit, x 2 / 3 , b. terminal cymes, x 3, c. flower, d. 

ditto in LS, e. stamen in dorsal and frontal view, f. receptacle containing ovary, disk, and style, all x 6, g. 

fruit, x 2 / 3 , h. CS of fruit, x 2 / 3 , i. embryo, x 6 (a-b BSIP 3080, c-f Clemens 1890, g-i BSIP 2809). 



1977] 



Cornaceae (Matthew) 



87 



median ridge inside, spreading or reflexed. Stamens 4-5(-6), or 8, alternating with 
the petals, erect in bud; when 8 in 2 alternate whorls of 4; filaments subulate, 
flattened; anthers cordate, dorsifixed, abutting on and alternating with the disk 
lobes, latrorse; connective ± protruding. Ovary inferior, turbinate, 1 -celled, sur- 
mounted by a prominent, fleshy, persistent disk c. V3 the height of the receptacle; 
invaginations of the disk abaxially 4-5 (fitting the filaments) and adaxially 8 or 10 
(fitting the thecae), becoming shallower with age; style stout, ribbed; stigma 
punctiform, sometimes deeply 2-fid or 4-5-lobed, lobes sometimes reflexed. Ovule 
1, pendulous laterally from the roof of the cell. Drupe subglobose to oblong, sur- 
mounted by calyx and disk; pericarp thin or thick, dark purple to blue when ripe; 
endocarp woody, sulcate on one side externally and internally deeply protruding 
into the fruit cavity as a wedge-shaped or swollen incomplete septum. Seed fitting 
the fruit cavity ; testa membranous; endosperm copious; embryo small; cotyledons 
foliaceous ; radicle elongate. 

Distr. About 13 spp. in SE. Asia (Western Ghats & Ceylon, NE. India, Bhutan, Burma, Thailand, 
Indo-China, S. Yunnan, Hainan) through Malesia to New Britain and the Solomon Islands. Fig. 2. 

Ecol. Primary and secondary forest, often in moist habitats, from sea-level up to 180O(-240O) m. 

Fossil endocarps of Mastixioids are found in quantity in the warmer Tertiary in Europe, Great Britain 
and North America. Cf. Kirchheimer, Die Laubgewach.se der Braunkohlzeit (1957) and D. H. Mai, 
Palaontol. Abhandl. Deut. 2 (1) (1964). The Pleistocene Glacial Epoch is held responsible for the contrac- 
tion of the range, similarly as happened to Symplocos, Meliosma, and so many other genera of the Tertiary 
mixed mesophytic forest on the northern hemisphere. 

Tax on. Mastixia was subdivided into two subgenera by Wangerin (1910) on the 4- and 5-merousness 
of the flowers respectively. Though this character is still used for discrimination of species, it seems 




Fig. 2. Range of the living species of the genus Mu\ti\iu Hi . For each district, island or island group the 
number of species is given, above the hyphen the endemic ones, below the hyphen the non-cndcmn. ones 

(occurring in more than one district). 



88 Flora Malesiana [ser. I, vol. 8 2 

artificial for subgeneric rank. Instead, I have proposed another subdivision (1976) into two subgenera, 
in one of which (subg. Manglesia) the stamens number 8 and are arranged into 2 whorls, while in subg. 
Mastixia the stamens number 4-5(-6) and stand in 1 whorl. Other differential characters support this 
subdivision ; see also the key. 

Anatomy. For general anatomical surveys also giving the older literature see Solereder, Syst. Anat. 
Dicot. Stuttgart (1899) 487-495 and ibid. (1908) 171-172; Metcalfe & Chalk, Anat. Dicot. Oxford 
(1950) 735-741. Additional selected references: Moll & Janssonius, Mikr. 3 (1918) 722-737 (wood 
anatomy); Adams, J. Elisha Mitchell Sci. Soc. 65 (1949) 218-244 (comparative wood anatomy); Jans- 
sonius, Blumea 6 (1950) 424 (wood anatomical affinities); Versteegh, Acta Bot. Need. 17 (1968) 151- 
159 (wood anatomy). 

The wood of Mastixia like that of most other Cornaceae is primitive. It has diffuse, exclusively solitary 
vessels with scalariform perforations (many-barred), fibre-tracheids, diffuse parenchyma, and hetero- 
geneous rays. Moll & Janssonius I.e. reported vertical intercellular canals in Mastixia rostrata and 
M. trichotoma. The latter are absent from M. tetrapetala studied in Leiden. The leaf and twig anatomy of 
Mastixia is characterized by the occurrence of secretory canals. This important feature is absent from the 
other genera of the Cornaceae. Their presence in Mastixia can be used as an argument to stress the 
affinities of Cornaceae with Araliaceae and Umbelliferae of the Cornales for which families they are 
typical. — P. Baas. 

Galls. Only two galls have been described by Docters van Leeuwen (Ned. Kruidk. Arch. 51, 1941, 
207) in the species where they most occur, viz M. rostrata and M. trichotoma, both caused by aphids. They 
occur, however, rather random in many species and varieties, with preponderance in ser. Oppositae. 
None have been found yet in species of subg. Manglesia. There are four kinds : on the stem, the leaf, the 
inflorescence, and the fruit. Sometimes they can be quite large, as has been cited under the species. See 

fig. 3. 

Uses. Although trees may reach a considerable size, the scattered occurrence does not contribute to 
general use as timber ; besides, the timber is not of good quality and is only used for minor purposes. Cf. 
Burkill, Diet. (1935) 1428. 

Notes. In key and descriptions the width of the submature flower is that of the corolla. 

About the use of the term 'merousness' of the flower it should be remarked that this cannot be used in 
the strict sense, as 4- and 5-merous flowers often occur in one inflorescence. If it is said 'basically 4-merous', 
this means that at least 80% of the flowers are 4-merous and the same holds for basically 5-merous 
flowers, so that the prevalent pattern is obvious. 

Moreover it should be remarked that the number of sepals frequently tends to be higher than that of 
petals and stamens. 

In exceptional cases identification of sterile or immature material must remain uncertain. 

Unfortunately no separate key can be provided for fruiting material. 

KEY TO THE SPECIES 

1. Stamens 8, in 2 whorls of 4. Inflorescence branches 4-angular (at least when young). Calyx subtruncate 
with minute, acute teeth. Bracts caducous. Pedicels of lateral flowers of terminal cymes over 5 mm, 
slender. Septum of endocarp swollen to at least Va of the diameter of the fruit. Branchlets subterete. 

Domatia occasional, suborbicular. Subg. Manglesia 1. M. octandra 

1. Stamens 4-5(-6), in one whorl. Inflorescence branches terete. Calyx distinctly lobed. Bracts subper- 

sistent. Septum of endocarp wedge-shaped. Subg. Mastixia. 
2. Inflorescence branches of the first order (sub)opposite or decussate. Branchlets and leaves generally 
(sub)opposite or decussate; nodes flattened. Fruits generally ovoid. Ser. Oppositae. 
3. Flowers basically 5-merous. 

4. Sepals less than half as long as wide. Inflorescence subglabrous to puberulous. Fruit ovoid to 
oblong, with inconspicuous persistent sepals 2. M. kaniensis 

4. Sepals almost as long as wide. Inflorescence velutinous to woolly. Fruit elongate-ovoid, with con- 
spicuous persistent sepals 3a. M. trichotoma var. korthalsiana 

3. Flowers basically 4-merous. 

5. Sepals almost as long as wide. Inflorescence puberulous to woolly. Corolla puberulous to villous 
outside. Leaves acute or shortly acuminate, 5-24 by 2-12 cm. Fruit with conspicuous persistent 
se p a l s 3. M. trichotoma 

5. Sepals less than half as long as wide. Inflorescence (sub)glabrous. Corolla glabrous outside. Leaves 

abruptly caudate to cuspidate, 4-12 by 2-5V 2 cm. Fruit with obscure calyx teeth. 
6. Leaves strictly opposite; petioles stout. Leaves thick-coriaceous; nervation prominent, with 
intermediary nerves. Inflorescence stout with lower bracts up to 5 mm. Fruit 1 1 / 2 cm 

4. M. eugenioides 
6. Leaves (sub)opposite or alternate ; petioles slender. Leaves chartaceous to subcoriaceous ; nerva- 
tion rather weak, without intermediary nerves. Inflorescence slender with bracts all under 3 mm. 

Fruit up to 1 cm 5. M. rostrata 

2. Inflorescence branches of the first order scattered. Branchlets and leaves scattered; nodes terete. 
Fruit generally ellipsoid or oblong. Ser. Alternae. 
7. Branchlets woolly. Leaves 13-30 by 5V 2 -15 cm, with midrib and nerves (even veinlets) woolly to 
villous; petioles stout, 4 cm or longer, woolly. Fruit over 4 by 2 cm. Flowers 5-merous 

6. M. macrocarpa 



1977] Cornaceae (Matthew) 89 

7. Branchlets not woolly. Petioles up to 4 cm. Fruit up to 4 cm long. 
8. Flowers basically 4-merous. 
9. Leaves glaucous and waxy below, thick-coriaceous, with intermediary nerves; apex apiculate. 

Sepals as long as wide 7. M. glauca 

9. Leaves not glaucous and waxy below, without intermediary nerves; apex other than apiculate. 
Calyx teeth at most half as long as wide. 
10. Leaves crowded at apices of branchlets, thick-coriaceous; acute to acuminate. Inflorescence 

branches stout, compact. Fruit ellipsoid, l 1 2 cm 8. M. tetrapetala 

10. Leaves evenly spread, chartaceous to subcoriaceous ; apex caudate (over 1 cm). Inflorescence 

branches rather slender 5. M. rostrata 

8. Flowers basically 5-merous. 
11. Leaves abruptly cuspidate (over 1 cm); nerves arcuate, clearly impressed above. Inflorescence 
raceme-like, seldom branched more than twice. Petals densely silky outside. Branchlets slender. 

Fruit oblong, l l ' 2 -2 by 0.8-1 cm 9. M. cuspidata 

11. Leaves other than abruptly cuspidate; nerves not arcuate but mostly sharply prominent, veins 
mostly distinct. Inflorescence usually branched twice or more, not terminating into a dichasium. 
Petals glabrous to appressed hairy. Fruit ovoid to oblong, 2 l U-VI 2 by 1-1 X U cm 

10. M. pentandra 

1. Subgenus Manglesia 

Matthew, Blumea 23 (1976) 64, f. 1 (map) & 2. 

Branchlets and leaves decussate. Stamens 8, in 2 whorls. Inflorescence branches 
4-angular. Calyx subtruncate. Fruit with swollen septum. 

Distr. 2 spp., in NE. India, N. Burma, NW. Thailand, Central Sumatra. 

1. Mastixia octandra Matthew, Blumea 23 under 3 mm, glabrous. Submature flower bud 
(1976) 65, f. 3 (map). 3 mm 0. Calyx subtruncate, thin; teeth 4, minute, 

Tree up to 25 m; d.b.h. up to 90 cm. Branchlets acute, thin. Petals 4, thick, glabrous outside, 

slender, decussate, terete, glabrous. Leaves decus- Stamens 8. Ovary glabrous. Fruit turbinate, 1 cm. 
sate, ovate to elliptic, 4-8 by l\' 2 -3 cm, chartaceous, Distr. Malesia: West Central Sumatra, once 

glabrous; base cuneate; apex acuminate; nerves found. 

6-8 pairs, with intermediary ones; veins distinct on Ecol. Mountain forest, 1700-1800 m. 
both surfaces; an occasional subcircular doma- Notes. Easily distinguished from the continen- 

tium at the axil of nerves; petiole 1-172 cm, ta ' Asian M. euonymoides Prain by smaller, 

slender. Inflorescence up to 15 cm, slender, glab- chartaceous leaves, suborbicular domatia, more 

rous, branched up to 5 times; branches of the first slender, lax and elongate inflorescence parts, the 

order decussate; pedicels of lateral flowers of thin calyx with acute teeth, and the generally 

terminal cymes over 5 mm, slender. Bracts ovate, pedicelled middle flower of the cymes. 

2. Subgenus Mastixia 

Cf. Matthew, Blumea 23 (1976) 66, f. 1 (map) & 2. — Mastixia subg. Tetra- 
mastixia et Pentamastixia Wangerin, Pfl. Reich Heft 41 4 (1910) 21, 25. 

Branchlets and leaves scattered or (sub)opposite. Stamens 4-5(-6), in one whorl. 
Inflorescence branches terete. Calyx lobed. Septum of the fruit wedge-shaped. 

Distr. 11 spp., covering the entire range of the genus. 

1. Series Oppositae ledermannii Melcm. Bot. Jahrb. 60 (1925) 173. — 

M. pentandra (non Bi.) Danser, Blumea I (1934) 
Matthew, Blumea 23 (1976) 66. 50, p.p.- Matthew, Blumea 23 (1976) 67. — Fig. 

Inflorescence branches of the first order (sub)- i ? 3 a ^b (galls). 

opposite or decussate. Branchlets and leaves ditto; 'Tree up to 31m; d.b.h. up to 75( 90) cm. 

nodes flattened. Emit usually ovoid. Branchlets stout or slender, (suh)opposite, sub- 

I )i st r. Throughout Malesia, in continental Asia glabrous to vclutinous. Leaves (sub)opposite, 

only in Pensinsular Thailand. elliptic, obovatc, oblong or oblanceolatc, (3 1 , > 

4' 2 ik by 2 B cm, chartaceous to thick coriaceous, 

2. \!;isti\i.i kaniemis Miimi. Mot. Jahrb. 60 sub^labious, rarely densely vclutinous; base 
M'V25) 172; DANKR, Blumea 1 (1934) 52. — M. attenuate to truncate ; apex acuminate to caudate; 



90 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 3. Galls of Mastixia. a. M. kaniensis Melch. spp. ledermannii (Melch.) Matthew, b. M. kaniensis 
Melch. ssp. kaniensis, c. M. pentandra Bl. ssp. chinensis (Merr.) Matthew, d-e. M. pentandra Bl. ssp. 
philippinensis (Wangerin) Matthew, f. M. rostrata Bl. ssp. caudatifolia (Merr.) Matthew, g. M. tricho- 
toma Bl. var. korthalsiana (Wangerin) Danser, h. M. trichotoma Bl. var. rhynchocarpa Danser. All nat. 
size, except g x 2 (a Clemens 5361, b Schlechter 17703, c Lace 5641, d Wenzel 1150, e FB 2201, 

/KOSTERMANS 12573, g KOSTERMANS 7316, h KOSTERMANS 7620). 



1977] 



Cornaceae (Matthew) 



91 



nerves 3— 9(— 1 1) pairs, sometimes arcuate; veins 
usually obscure; petiole l-2(-4)cm, stout or 
slender. Inflorescence up to 8 cm, stout or slender, 
subglabrous to puberulous, branched up to 3-(4) 
times, at times terminating in a dichasium, branches 
of the first order (sub)opposite. Bracts triangular 
to lanceolate, up to 4 mm, puberulous to veluti- 
nous. Submature flower bud \ l l 2 -2 x l 2 mm 0. 
Sepals (4-)5(-6-7), broader than long. Petals 
(4-)5(-6), thick or thin, glabrous or puberulous 
outside. Stamens (4-)5(-6). Ovary sparsely puberu- 
lous. Fruit ovoid to oblong, l 1 j-2 1 t by 1-1 ' 2 cm, 
dull or shining when dry; persistent disk incon- 
spicuous to prominent ; sepals inconspicuous. 

Distr. Malesia: Moluccas, New Guinea, New 
Britain, and the Solomon Islands. Fig. 4. 

Note. Two replacing subspecies are distin- 
guished. The maximum degree of fluctuation in the 
number of flower parts occurs in the New Guinea 
— Solomons area. 



KEY TO THE SUBSPECIES 



1. Leaves chartaceous to subcoriaceous ; branch- 
lets and inflorescence axes slender; petals thin, 
glabrous outside a. ssp. kaniensis 

1 . Leaves thin- to thick-coriaceous ; branchlets and 
inflorescence axes usually stout; petals thick, 
puberulous outside. ... b. ssp. ledermannii 

a. ssp. kaniensis. — Fig. 1, 3b (galls). 

Branchlets slender, subglabrous to velutinous. 
leaves (i l l 2 -)4 l l 2 -l4 l l 2 by 2-6 cm, chartaceous to 
subcoriaceous; base attenuate to cuneate; nerves 
3-7 pairs, sometimes arcuate, prominulous below; 
petiole 1-2 cm. Inflorescence up to 6 cm, slender, 
branched 2(-3) times, lax. Flowers relatively small. 
Bracts triangular to lanceolate, under 3 mm. Sub- 
mature bud l'/ 2 mm 0. Petals (4-)5, relatively 



thin, glabrous outside. Stamens (4-)5. Fruit ovoid 
to oblong, 2-2Va by 1 V4-IV2 cm. 

Distr. Malesia: East New Guinea, New Britain, 
and Solomon Islands. Fig. 4. 

Ecol. Common in primary forests from low 
altitude up to 1200 m. Fl. fr. Jan.-Dec. Leaf- and 
fruit-galls occur. 

Note. Occasional specimens in New Guinea are 
densely velutinous, others are less so. The basal 
branches of the inflorescence of the first order are 
at times subtended by foliage leaves. 

b. spp. ledermannii (Melch.) Matthew, Blumea 23 
(1976) 67. — M. ledermannii Melch. — Fig. 3a 
(galls). 

Branchlets stout, often rusty puberulous when 
young, subglabrous later. Leaves 6-18 by 2V 2 -8 
cm, thin- to thick-coriaceous; base attenuate to 
truncate; nerves 3-9(-l 1) pairs, sometimes arcuate, 
obscure to prominent below ; petiole 1 l l 2 -2(-4) cm. 
Inflorescence up to 8 cm, stout, branched 3-(4) 
times, compact. Flowers relatively large. Bracts 
lanceolate below, up to 4 mm. Submature flower 
bud 2 l l 2 mm 0. Petals (4-)5(-6), thick, puberulous 
outside. Stamens (4-)5(-6). Fruit ovoid, IVj- 
2V 2 by l-lV4cm. 

Distr. Malesia: Moluccas and New Guinea. 
Fig. 4. 

Ecol. Primary and secondary forests, 100-1800 
m. Fl. fr. Jan.-Dec. Leaf-galls occur. 

Vern. New Guinea: bie, bon, Muju, labak- 
kobilien, Mooi, masjiw, Wandammen, samuwin, 
Biak. 

Notes. In some specimens a dense indumentum 
is found. 

Though the two subspecies are clearly replacing, 
some specimens of ssp. ledermanii occur in the area 
occupied by ssp. kaniensis, but at higher altitude 
than ssp. kaniensis in this area. 




Fig. 4. Localities of three species and two subspecies of Mastixia. 



92 



Flora Malesiana 



[ser. I, vol. 8 2 



3. Mastixia trichotoma Bl. Bijdr. (1826) 655; DC 
Prod. 4 (1830) 275; Bl. Mus. Bot. 1 (1850) 257 
f. 58; Miq. Fl. Ind. Bat. 1, 1 (1856) 772, incl. var 
laxa Miq. I.e.; K. & V. Bijdr. 5 (1900) 90; Wan 
gerin, Pfl. Reich Heft 41 4 (1910) 24, f. 1A-E 
Koord. Atlas 1 (1913) t. 190; Fl. Tjib. 2 (1923) 
Danser, Blumea 1 (1934) 57, incl. var. tenuis 
acuminatissima, clarkeana, korthalsiana, maingayi 
benculuana, rhynchocarpa et simalurana Danser 
I.e. 61-65; Back. & Bakh./. Fl. Java 2 (1965) 159 
Meijer, Bot. News Bull. Sandakan 8 (1967) 65 
ibid. 10 (1968) 179, illus.; Matthew, Blumea 23 
(1976) 68. — M. laxa Bl. Mus. Bot. 1 (1850) 257, 
incl. var. angustifolia Bl. I.e. ; Wangerin, Pfl. 
Reich Heft 41 4 (1910) 24. — M. acuminatissima 
Bl. Mus. Bot. 1 (1850) 258; Miq. Fl. Ind. Bat. 1, 1 
(1856) 772, (1858) 1095; Wangerin, Pfl. Reich 
Heft 41 4 (1910) 22, f. IF. — M. caesia Bl. Mus. 
Bot. 1 (1850) 258. — M. kimanilla Bl. I.e. 258; 
Miq. Fl. Ind. Bat. 1, 1 (1856) 772, (1858) 1095, 
incl. var. caesia Miq. I.e. 772; K. & V. Bijdr. 5 
(1910) 94; Wangerin, Pfl. Reich Heft 41 4 (1910) 
25. — M. maingayi Clarke, Fl. Br. Ind. 2 (1879) 
746; King, J. As. Soc. Beng. 71, ii (1902) 74, incl. 
var. subtomentosa King, I.e. 75; Wangerin, 
Pfl. Reich Heft 41 4 (1910) 22. — M. junghuhniana 
(non Miq.) Clarke, Fl. Br. Ind. 2 (1879) 746. — M. 
clarkeana King, J. As. Soc. Beng. 71, ii (1902) 75 & 
var. macrophylla King, I.e. ; Wangerin, Pfl. Reich 
Heft 41 4 (1910) 24; Hall./. Beih. Bot. Centralbl. 
34, 2 (1916) 40. — M. korthalsiana Wangerin in 
Fedde, Rep. 4 (1907) 335, incl. var. macrophylla 
Wangerin, I.e. 336 et Pfl. Reich Heft 41 4 (1910) 
25, 26; Hall./. Beih. Bot. Centralbl. 34, 2 (1916) 
40. — M. propinqua Ridl. J. Fed. Mai. St. Mus. 4, 
1 (1909) 25; Fl. Mai. Pen. 1 (1922) 890. — Vitex 
premnoides Elmer, Leafl. Philip. Bot. 8 (1915) 2874. 
— M. premnoides Hall. /. Beih. Bot. Centralbl. 
34, 2 (1916) 41 ; Merr. Philip. J. Sc. 13 (1918) 43; 
En. Philip. 3 (1923) 242. — M. rostrata {non Bl.) 
Ridl. Fl. Mai. Pen. 1 (1922) 890. — Fig.3g-h (galls). 

Tree up to 40 m; d.b.h. up to 50(-150)cm; 
branchlets stout or slender, opposite, puberulous 
to woolly. Leaves opposite, ovate, elliptic to 
oblong, 5-24(-28) by 2-12 cm, thin- to thick- 
coriaceous, subglabrous to villous below; base 
cuneate, obtuse or attenuate; apex acute to acu- 
minate; nerves 5-15 pairs, impressed above, 
prominulous to prominent below, at times arcuate; 
veins prominulous to prominent below; petiole 
V U-2 1 1 2 {-V 1 2 ) cm, stout or slender. Inflorescence 
up to 15 cm, stout or slender, compact or lax, 
puberulous to woolly; branched up to 5 (or 6) 
times; branches of the first order opposite; higher 
order bracts triangular, more or less connate, 
villous to woolly; lower bracts lanceolate, up to 
10 mm, villous to woolly. Submature flower bud 
2-3V2 mm 0- Sepals 4 or 5, as long as wide, thick, 
puberulous to villous. Petals 4 or 5, thick, puberu- 
lous to villous outside. Stamens 4 or 5. Ovary 
puberulous to villous. Fruit ovoid to elongate, 
acute, IV2-3V4 by V2-2cm; persistent disk incon- 
spicuous to prominent; sepals prominent. 

Distr. Peninsular Thailand and throughout 
Malesia, except New Guinea. Fig. 5. 

Notes. Danser (I.e. 59-61) adequately dis- 
cussed variations within the species; most of the 
vernacular names he listed (I.e. 72-73) belong to the 
present species. 







1 



Fig. 5. Localities of Mastixia trichotoma Bl. and 
its varieties. 



Figure 1 of Wangerin (I.e. 23) is rather sche- 
matic. The position of ovule attachment and the 
length of the sepals are inaccurately drawn. 

KEY TO THE VARIETIES 

1. Inflorescence villous to woolly. 
2. Twigs woolly. Leaves 9-20 cm long, thick- 
coriaceous, villous to woolly; nerves often 
arcuate; base obtuse to truncate. Fruit ovoid, 
more than 1 1 / 2 cm . . . . c. var. maingayi 
2. Twigs not woolly. Leaves 5-15 cm long, 
chartaceous to subcoriaceous, subglabrous to 
puberulous; nerves seldom arcuate; base 
attenuate to cuneate. Fruit elongate-ovoid, up 
to l'/icm 0. 
3. Inflorescence compact. Sepals, petals, and 

stamens 4 a. var. trichotoma 

3. Inflorescence very lax. Sepals, petals, and 

stamens 5 b. var. Korthalsiana 

1. Inflorescence subglabrous to puberulous. 
4. Fruit with prominent persistent disk, over 2V2 
by IV2 cm. Leaves 10-24 by 5-12 cm. Inflores- 
cence robust, up to 1 5 cm, branched 5 (to 6) 
times, not terminating in a dichasium. 

d. var. rhynchocarpa 

4. Fruit without prominent persistent disk, up to 

2 by 1 cm. Leaves usually 5-12 by 2-4V2 cm. 

Inflorescence slender, up to 10 cm, branched 

3(-4) times, often terminating in a dichasium 

e. var. clarkeana 

a. var. trichotoma — M. laxa Bl. — M. trichotoma 
Bl. var. laxa Miq. — M. laxa Bl. var. angustifolia 
Bl. — M. acuminatissima Bl. — M. trichotoma 
Bl. var. acuminatissima Danser — M. caesia Bl. 
— M. kimanilla Bl. incl. var. caesia Miq. 

Branchlets rather slender, yellowish, puberulous 
to velutinous. Leaves elliptic to oblong, 5-15 by 
3-8 cm, chartaceous to subcoriaceous, sub- 
glabrous to puberulous ; base cuneate to attenuate ; 
apex acute to acuminate; nerves 5-8 pairs, promi- 
nulous to prominent below, seldom arcuate; 
petiole l'/4-2(-2V2) cm, rather slender. Inflores- 
cence up to 8 cm, compact, branched 3(-4) times, 
villous to woolly; basal bracts under 5 mm; 
terminal bracts often deeply boat-shaped. Sepals 4. 
Petals 4. Stamens 4. Fruit elongate-ovoid, 1 1 /2~3 by 



1977] 



Cornaceae (Matthew) 



93 



l-lV2cm; persistent disk inconspicuous; sepals 
prominent. 

Distr. Malesia: N. Sumatra, W.-E. Java, 
Borneo, Lesser Sunda Is. (Bali). Fig. 5. 

Ecol. In primary forest from low altitude up to 
1800 m. Fl. May-Jan., fr. July-March. Galls occur 
on stem and fruit. 

Vern. Java: djeret, hunt hiris, h. minjak, kendu, 
kibintili, kibunting, kidedak, kilumlum, kilun glum, 
kiminjan, kitenjo, (huru) mehmal, memah, palaglar 
minjak, tinggau, tenju. 

b. var. korthalsiana (Wangerin) Danser, Blumea 1 
(1934) 63; Matthew, Blumea 23 (1976) 70. — 
M. korthalsiana Wangerin. — Fig. 3g (galls). 

Branchlets rather slender, yellowish, subglab- 
rous to velutinous. Leaves subovate-elliptic, 10-12 
by 3-5 cm, subcoriaceous ; base long attenuate; 
apex acuminate; nerves 5-6 pairs, seldom arcuate; 
petiole l 1 4 -l 3 4 cm, rather slender. Inflorescence 
up to 10 cm, very lax, branched 3(^t) times, few- 
flowered, velutinous to woolly; bracts under 3 mm. 
Sepals 5. Petals 5. Stamens 5. Fruit elongate-ovoid, 
2 1 2 -3 by 1-1 l : cm, persistent disk inconspicuous, 
sepals prominent. 

Distr. Malesia: Sumatra, Borneo. Fig. 5. 

Ecol. In primary forest from low altitude up to 
640 m, often scattered. Rather prone to galls. 

Vern. Sumatra: medang kladi. E. Borneo: 
sirgam pipit, Sangkulirang I. 

Note. The lax and few flowered inflorescence, 
the 5-merous flowers, and elongate fruit distinguish 
this variety from var. trichotoma to which it is 
closely allied. 

c. var. maingavi (Clarke) Danser, Blumea 1 
(1934) 63; Matthew, Blumea 23 (1976) 70. — M. 
maingayi Clarke, incl. var. subtomentosa King — 
M.junghuhniana {non MlQ.) Clarke — M. rostrata 
(non Bl.) Ridl. — M. propinqua Ridl. 

Branchlets very stout, yellowish, woolly. Leaves 
ovate to elliptic, 9-20 by 4-1 1 cm, thick coriaceous, 
tough and stiff; base obtuse to truncate; apex acute 
to acuminate; nerves 5-6 pairs, deeply impressed 
above, very prominent below, often arcuate, 
numerous parallel intermediary veins conspicuous; 
petiole l l 2 -2' 2 cm, stout, villous to woolly. 
Inflorescence up to 15 cm, compact, branched 
4(-5) times, velutinous to golden woolly; basal 
bracts up to 1 cm, persistent. Sepals 4. Petals 4. 
Stamens 4. Fruit ovoid, 2 I / 2 -3 l /2 by I7r-2cm; 
persistent disk inconspicuous, sepals prominent. 

Distr. Malesia: Sumatra, Malay Peninsula (also 
Penang), Banka, Borneo. Fig. 5. 

Ecol. In primary forests with Dipterocarps;also 
in secondary or marsh forests, from the lowland to 
1400 m. Fl. Febr.-Aug. (Nov.), fr. April-Dec. Galls 
occur on leaves and fruit. 

Vern. Malaya: karu nuri, kayu btngkal bukit, 
kayu maura, medang. Banka: midang pttntung, 
m. puslr. Borneo : medang kanigara. 

Note. Easily recognized by the woolly indumen- 
tum, large and stiff leaves with prominent veins, 
and the numerous massive fruits. 

d. var. rhynchocarpa DANSER, Blumea 1 (1934) 64; 
Matthiw, Hlumca 23 (1976) 71. - M. trichotoma 
Bl. var. bvm uluana ct var. simalurana Dansi | 

<h (galls). 



Branchlets stout, dark, subglabrous. Leaves 
elliptic to oblong, 10-24(-28) by 5-12 cm, 
coriaceous, tough and stiff; base cuneate to obtuse; 
apex acute, acuminate to caudate; nerves 8-15 
pairs, very prominent below, seldom arcuate; 
petiole l 1 2-2 1 2 (-3 l / 2 ) cm, stout. Inflorescence up 
to 15 cm, compact, profusely branched up to 
5(-8) times, not terminating in a dichasium, sub- 
glabrous to puberulous; basal bracts up to 5 mm. 
Sepals 4(-5). Petals 4. Stamens 4. Fruit ovoid to 
elongate-ovoid, 2\' 2 -3 1 4 by lV 2 -2cm; persistent 
disk prominent, bulging; sepals prominent. 

Distr. Malesia: Sumatra (incl. Simalur I.), W. 
Java, Borneo, NE. Celebes, Moluccas (Ambon, 
Ceram). Fig. 5. 

Ecol. Common in primary lowland and mossy 
forest, up to 1800 m. Fl. April-Aug., fr. June- 
March. 

Galls. This variety is very prone to fruit galls 
and the largest ones in the genus (over \ x j 2 cm 0) 
occur here. 

Vern. Sumatra: aheldt, awa ahelat uding, awa 
enti, awa simangurach, tutun simangurah, Simalur 
I.; bung, medang tima, tanah, Bencoolen. Borneo: 
medang aima. Moluccas: soya. 

Note. This variety is noted for the generally 
large dimensions of leaves and inflorescence, 
though there is a reduction in size from Borneo to 
Moluccas. 

e. var. clarkeana (King) Danser, Blumea 1 (1934) 
62; Matthew, Blumea 23 (1976) 72. — M. 
clarkeana King, incl. var. macrophylla King — 
M. korthalsiana Wangerin var. macrophylla 
Wangerin — Vitex premnoides Elmer — M. prem- 
noides Hall. /. — M. trichotoma Bl. var. tenuis 
Danser. 

Branchlets slender, grey, subglabrous. Leaves 
oblong to elliptic-oblong, 5-12(-18) by 2-4(-8'/ 2 ) 
cm, coriaceous; base cuneate to obtuse; apex acute 
to acuminate; nerves 5-7 pairs, seldom arcuate; 
petiole l-l\' 2 cm, slender. Inflorescence up to 
10 cm, compact, branched 3(-4) times, often ter- 
minating in a dichasium. Sepals 4. Petals 4. 
Stamens 4. Fruit ovoid, VI 2 -2 by x l 2 -\ cm; per- 
sistent disk inconspicuous; sepals prominent. 

Distr. Peninsular Thailand (Pattani) and 
Malesia: Sumatra, Banka, Malay Peninsula, 
Borneo, Philippines (Mindanao). Fig. 5. 

Ecol. Primary forest, from low altitude to 
1 100 m. Fl. Jan. -Aug., fr. July-Febr. Galls occur on 
stem and fruit. 

Vern. Philippines: lamog. 

4. Mastixia eugenioides Matthew, Blumea 23 

(1976) 73. 

Tree up to 30 m ; d.b.h. up to 30 cm ; branchlets 
stout, opposite, glabrous. Leaves opposite, elliptic 
to oblong-elliptic, 4-12 by 2-5'/ 2 cm, thick- 
coriaceous, glabrous; base cuneate; apex acumi- 
nate to caudate; nerves 5-7 pairs, prominent 
beneath, with intermediary ones and distinct veins; 
petiole 1 '/ 2 -2'/j cm, stout. Inflorescence up to 8 cm, 
rather stout and compact, glabrous, up to 4 times 
branched, at times terminating in a dichasium : 
branches of the first order opposite; higher order 
bracts triangular, under J mm; lower bracts 
lanceolate, up to 5 mm, all glabrous. Suhmaturc 
flower bud 2 mm 0. Sepals 4, broader than long, 



94 



Flora Malesiana 



[ser. I, vol. 8 2 



thick, glabrous. Petals 4, thin, glabrous outside. 
Stamens 4. Ovary glabrous. Fruit (unripe) ovoid, 
2V2 by l'/2cm; persistent disk and sepals incon- 
spicuous. 

Distr. Malesia: Borneo (Sarawak, Brunei). 
Fig. 4. 

Ecol. Primary (often Dipterocarp) lowland 
forests, up to 400 m.Fl. July-Aug.,/r. Sept. 

Notes. Leaf scars conspicuous; inflorescence 
notably erecto-patent when young, spreading later. 
The inflorescence and flowers somewhat resemble 
those of M. rostrata ssp. rostrata, but the stout 
branchlets with strictly opposite leaves and stout 
petiole, prominent intermediary veins, and fruits of 
different shape with thick pericarp, make this 
species quite distinct. 

All the 9 collections are from a restricted area. 

5. Mastixia rostrata Bl. Mus. Bot. 1 (1850) 258; 
Miq. Fl. Ind. Bat. 1, 1 (1856) 773, (1858) 1095; 
K. & V. Bijdr. 5 (1900) 92; Wangerin, Pfl. Reich 
Heft 41 4 (1910) 22; Koord. Atlas 1 (1913) t. 191; 
Danser, Blumea 1 (1934) 52; Burk. Diet. (1935) 
1428; Doct.v.Leeuwen, Ned. Kruidk. Arch. 51 
(1941) 207; Back. & Bakh. /. Fl. Java 2 (1965) 
159; Meuer, Bot. News Bull. Sandakan 8 (1976) 
65; Matthew, Blumea 23 (1976) 73. — M. jung- 
huhniana Miq. Fl. Ind. Bat. 1, 1 (1856) 772. — 
M. margarethae Wangerin in Fedde, Rep. 4 (1907) 
335; Pfl. Reich Heft 41 4 (1910) 21. — M. cuspidata 
Bl. var. margarethae Hall./. Beih. Bot. Centralbl. 
34, 2 (1916) 41. — M. caudatifolia Merr. PI. Elm. 
Born. (1929) 233. 

Tree up to 30 m; d.b.h. up to 50 cm; branchlets 
slender, (sub)opposite or scattered, glabrous. 
Leaves (sub)opposite or scattered, elliptic to oblong- 
elliptic, 4-10 by 2-5 cm, chartaceous to sub- 
coriaceous, glabrous; base cuneate; apex caudate 
over 1 cm; nerves 4-6 pairs, prominulous beneath; 
veins obscure; petiole l-2 1; 2 cm, slender. Inflores- 
cence up to 6 cm, slender, compact or lax, sub- 
glabrous, up to 4 times branched, branches of the 
first order (sub)opposite or scattered; bracts 
triangular, under 3 mm, glabrous. Submature 
flower bud I-2V2 mm 0. Sepals 4, broader than 
long, thin, glabrous. Petals 4, glabrous outside. 
Stamens 4. Ovary glabrous. Fruit ovoid to oblong, 
IV2-3 by 3 / 4 -l cm, persistent disk prominent or 
inconspicuous, sepals inconspicuous. 

Distr. Malesia: Sumatra, Banka, Java, Borneo, 
Lesser Sunda Is. (Sumbawa, Flores). Fig. 4. 



KEY TO THE SUBSPECIES 

1. Submature flower bud 2\' 2 mm 0. Inflorescence 
compact, branches of the first o - der (sub)- 
opposite. Leaves (sub)opposite. Galls absent 

a. ssp. rostrata 

1. Submature flower bud 1 mm 0. Inflorescence 
lax, branches of the first order scattered. Leaves 
scattered. Galls frequent. . b. ssp. caudatifolia 

a. ssp. rostrata — M. junghuhniana Miq. 

Branchlets (sub)opposite. Leaves (sub)opposite, 
less often scattered ; petiole 1 1 / 2 -2 cm. Inflorescence 
compact, generally under 4 cm, glabrous; primary 
branches (sub)opposite. Submature flower bud 
2 1 i ' 2 mm 0. 



Distr. Malesia: West & Central Java, Lesser 
Sunda Is. (Sumbawa, Flores). Fig. 4. 

Ecol. Moist forest, from low altitude up to 
1400 m. Fl. March, very fragrant, fr. May-July. 
Galls absent. Ripe fruit dark blue. 

Vern. Java: daun kaju tendjo, daun kitadjas, 
huru gading, kilburoy, kilejas, kitendjo, kitindjo, 
lalakina, tjangkar. Flores: bumis, rau, tapaaeke. 

Notes. Specimens from the Lesser Sunda Is. 
have larger leaves than those from Java. 

The reference in Kanjilal & Das (Fl. Assam 2, 
1938, 371) to this species seems erroneous, as it 
does not occur on the Asian continent. If the state- 
ment "stamens 3" is correct, the plant can even not 
belong to Mastixia. 

b. ssp. caudatifolia (Merr.) Matthew, Blumea 23 
(1976) 74. — M. margarethae Wangerin — M. 
caudatifolia Merr. — Fig. 3f (galls). 

Branchlets scattered. Leaves scattered; petiole 
up to lVzCm. Inflorescence very lax, up to 6 cm, 
puberulous at the nodes; primary branches 
scattered. Submature flower bud 1 mm 0. 

Distr. Malesia: northern half of Sumatra, 
Banka, Borneo. Fig. 4. 

Ecol. Primary forest, from the lowland up to 
1 600 m. Fl. June-Oct., fr. Aug.-March. Globose to 
elongate galls are common, specially those on 
fruits. Sometimes they resemble a legume and can 
be up to 3V 2 cm long. 

Vern. Borneo: patoli entelit, Iban lang. 

Note. It is almost impossible to separate sterile 
materials of ssp. caudatifolia from those of M. cus- 
pidata, though their flowers are entirely different. 

2. Series Alternae 

Matthew, Blumea 23 (1976) 75. 

Inflorescence branches of the first order scat- 
tered; branchlets and leaves scattered; nodes 
terete; fruit generally ellipsoid or oblong. 

Distr. Ceylon and continental Asia; through 
Malesia, but absent in New Guinea and Solomon 
Is. 

6. Mastixia macrocarpa Matthew, Blumea 23 
(1976) 75. 

Tree up to 21 m; d.b.h. up to 20 cm; branchlets 
stout, scattered, woolly. Leaves scattered, elliptic- 
oblong to oblong, 13-30 by 5V 2 -15cm, subcoria- 
ceous, villous, especially below; base cuneate, at 
times slightly oblique; apex acuminate; nerves 
7-10 pairs, prominent below, villous; veins promi- 
nulous, puberulous to villous; petiole 4-7 cm, 
stout, woolly. Inflorescence up to 9 cm, stout, 
woolly, branched up to 4 times; branches of the 
first order scattered ; bracts triangular to lanceolate, 
up to 8 mm, densely woolly. Submature flower bud 

4 mm 0. Sepals 5, broader than long, villous. 
Petals 5, velvety outside. Stamens 5. Ovary densely 
villous. Fruit oblong-ovoid, 4-4 1 / 2 by 2 cm; per- 
sistent disk inconspicuous, sepals prominent, up to 

5 mm. 

Distr. Malesia: Borneo (Sarawak), Philippines 
(Luzon); 2 collections. Fig. 6. 

Ecol. Lowland forest. Fl. Oct., fr. June. Sticky 
resin on the branches; fruits very pale green. 

Note. Leaves and fruits are the largest known in 
the genus ; lenticels up to 3 by 1 mm ; leaf scars up 



1977] 



Cornaceae (Matthew) 



95 



to 4 by 4 mm ; peduncles of terminal cymes up to 
6 mm ; pedicels 2 mm ; sepals 2 mm broad at the 
base; filaments 3 mm; anthers l 1 < 2 mm; receptacle 
3-3V2 by 2 mm; style 4 mm; stigma 5-lobed, 
appearing bifid. 




# n. cq*pld«t* »1« 






QC^S-£^^ 3 ^^> 



Fig. 6. Localities of three species of Mastixia. 

7. Mastixia glauca Matthew, Blumea 23 (1976) 
76. 

Tree up to 15 m; branchlets stout, scattered, 
glabrous. Leaves scattered, obovate, 7-16 by 
4 l 2 -8' 2 cm, thick-coriaceous, glaucous and waxy 
below, glabrous; base obtuse; apex apiculate; 
nerves 4-5 pairs, with intermediary ones, all 
obscure; veins obscure; petiole 2-3 1 2 cm, stout, 
glabrous. Inflorescence up to 5 cm, rather stout and 
compact, subglabrous to sparsely puberulous, 
branched up to 5 times ; branches of the first order 
scattered; bracts triangular, under 3 mm. Sub- 
mature flower bud 3 mm 0. Sepals 4(-5), as long 
as broad, sparsely puberulous. Petals 4(-5), thick, 
appressed-hairy outside. Stamens 4(-5). Ovary 
sparsely puberulous. 

Distr. Malesia: Borneo (Sarawak: Mt Santu- 
bong), 3 collections Fig. 6. 

Ecol. Lowland forest. Fl. April-May. 

Note. Tender bark of branchlets yellowish; 
inflorescence clearly broader than long; 5-merous 
flowers only occasional. 

8. Mastixia tetrapetala Mirk. Philip. J. Sc. 13 
(1918) Bot. 42; En. Philip. 3 (1923) 242; Matthew, 
Blumea 23 (1976) 76, f. 5 (map). — M. pachvphylla 
Merr. Philip. J. Sc. 13(1918) Bot. 325; En. Philip. 
3 (1923) 241. — M. crassifolia Merr. Philip. J. Sc. 
26(1925)486. 

Tree up to 8(-15)m; branchlets very stout, 
scattered to subopposite, subglabrous. Leaves 
scattered to subopposite, crowded at apices of 
branchlets, obovate to oblanceolate, 5-15 by 
2-7 cm, thick coriaceous, glabrous; base cuneate; 
apex acute to acuminate; nerves 6 8(12) pairs, 
usually prominent below; veins prominulous 
below; petiole l-2',cm, stout, glabrous. In- 
• nee up to 3 (6) cm, stout, very compact, 
puberulous to villous, 2( ~\) times branched; 
branches of the first order scattered; higher order 
bracts triangular, under 3 mm; lower ones lanceo- 
late up to 5 mm, puberulous. Submature flower 



bud 5 mm 0. Sepals 4, broader than long, glabrous 
to appressed-hairy. Petals 4, thick, glabrous to 
appressed-hairy. Stamens 4. Ovary glabrous to 
appressed-hairy. Fruit ellipsoid, 2-3 by l 1 /*- 
lVa cm; persistent disk and sepals inconspicuous. 

Distr. Malesia: Philippines (Luzon, Catan- 
duanes), 7 collections. 

Ecol. Primary forest, from low altitude up to 
2300 m. Fl. Febr.-March, fr. Sept.-Febr. 

Notes. Branchlets stout with conspicuous leaf 
scars and fibrous bark. Phyllotaxis tends to be 
obscured owing to congestion of parts: leaves are 
generally crowded towards the apices of branch- 
lets. Inflorescence branches do not always elongate 
as in other species. 

The species is quite distinct and stands rather 
isolated from others in the stoutness of parts, the 
large, 4-merous flowers, and the large, ellipsoid 
fruits. 

Two other species of Merrill, M. pachyphylla 
and M. crassifolia are considered conspecific with 
M. tetrapetala. There are indeed certain differences : 
leaves of M. crassifolia generally have 8-12 nerves 
per side prominent below, and massive fruits. 
M. pachyphylla has (sub)opposite leaves and pri- 
mary inflorescence branches. However, when 
examined together, it is seen that both M. pachy- 
phylla and M. crassifolia are extreme variations of 
M. tetrapetala. 

9. Mastixia cuspidata Bl. Mus. Bot. 1 (1850) 256; 
Miq. Fl. Ind. Bat. 1, 1 (1856) 772; Hall./. Beih. 
Bot. Centralbl. 34, 2 (1916) 41 ; Danser, Blumea 1 
(1934) 55, excl. var. margarethae; Matthew, 
Blumea 23 (1976) 79. — M. pentandra Bl. var. 
cuspidata Miq. Fl. Ind. Bat. 1, 1 (1858) 1095; 
Wangerin, Pfl. Reich Heft 41 4 (1910) 26. — M. 
bracteata Clarke, Fl. Br. Ind. 2 (1879) 746; King, 
J. As. Soc. Beng. 71, ii (1902) 73; Wangerin, 
Pfl. Reich Heft 41 4 (1910) 26, f. 1 G-K, N-O; 
Danser, Blumea 1 (1934) 54. 

Tree up to 24 m; d.b.h. up to 40 cm; branchlets 
very slender, scattered, subglabrous. Leaves 
scattered, obovate, elliptic or oblong, 4— 12(-I6) by 
2-4(-6) cm, subcoriaceous, glabrous; base cuneate; 
apex abruptly cuspidate (over 1 cm), oblique; 
nerves 4 (or 5) pairs, arcuate, impressed above; 
veins obscure; petiole 1-1 '/ 4 cm, slender. Inflores- 
cence up to 4 cm, rather slender, subglabrous to 
puberulous, 2(-3) times branched; branches of the 
first order scattered ; higher order bracts subulate, 
lower ones foliaceous, over 10 mm, passing into 
foliage leaves. Submature flower bud 3 mm 0. 
Sepals 5, broader than long, subglabrous. Petals 
5, thick, densely appressed-hairy outside. Stamens 
5. Ovary densely appressed silky-hairy. Fruit 
oblong, 1Vj-2 by 3 / 4 -l cm; persistent disk and 
sepals inconspicuous. 

Distr. Malesia: Sumatra, Banka, Malay 
Pensinula, Borneo. Fig. 6. 

Ecol. Primary and secondary forests, from low 
altitude up to 900 m. 

Vern. Sumatra: bebung, kundur. Banka: min- 
kapm, Malay Peninsula: dadaru. Borneo: biansu- 
gunong, Sarawak. 

10, Mastixia pentandra Bi . Bijdr. (1826) 654; DC. 
Prod. 4 (1830) 275; Bl. Mus. Bot. I (1850) 256; 
MlQ. FL Ind. Bat. 1, 1 (1856) 771, (1858) 1095; 



96 



Flora Malesiana 



[ser. I, vol. 8 2 



K. & V. Bijdr. 5 (1900) 88; Merr. Philip. J. Sc. 1 
(1906) Suppl. Ill; Danser, Blumea 1 (1934) 49; 
Back. & Bakh./. Fl. Java 2 (1965) 159; Matthew, 
Blumea 23 (1976) 80, f. 5 (map), 6. — M. arborea 
[non (Wight) Bedd.] Clarke, Fl. Br. Ind. 2 (1879) 
745, p.p.; Kanjilal & Das, Fl. Assam 2 (1938) 
370; Hundley & Chit, Trees Shr. Burma ed. 3 
(1961) 119. — M. cambodiana Pierre, Fl. Coch. 
(1892) t. 260 B; Wangerin, Pfl. Reich Heft 41 4 
(1910) 29; Evrard, Fl. Gen. I.-C. 2(1923) 1 195. — 
M. scortechinii King, J. As. Soc. Beng. 71, ii 
(1902) 73; Wangerin, Pfl. Reich Heft 41 4 (1910) 
27, f. 1 L-M; Danser, Blumea 1 (1934) 56. — M. 
philippinensis Wangerin in Fedde, Rep. 10 (1912) 
273; Merr. En. Philip. 3 (1923) 241; Chao, 
Taiwania 5 (1954) 94, 99, f. 37; Li & Chao, Quart. 
J. Taiwan Mus. 7 (1954) 124, f. 19. — M. parvi- 
folia Hall./. Beih. Bot. Centralbl. 34, 2 (1916) 41 ; 
Danser, Blumea 1 (1934) 51. — M. subcaudata 
Merr. Philip. J. Sc. 13 (1918) 43; En. Philip. 3 
(1923) 242. — M. megacarpa Ridl. Fl. Mai. Pen. 1 
(1922) 891. — M. chinensis Merr. Sunyatsenia 3 
(1937) 256; Li, Taiwania 1 (1938) 94. — M. alterni- 
folia Merr. & Chun, Sunyatsenia 5 (1940) 153. — 
M. poilanei Tardieu, Fl. Camb. Laos & Vietnam 
8 (1968) 16. — Fig. 3c-e (galls). 

Tree up to 37 m; d.b.h. up to 75 cm; branchlets 
slender or stout, scattered, subglabrous to puberu- 
lous. Leaves scattered, obovate, elliptic to oblong- 
elliptic, 4-16 by 2-8 cm, chartaceous to thick- 
coriaceous, subglabrous : base cuneate to attenuate ; 
apex acute or acuminate to caudate; nerves 4-7(-9) 
pairs, distinct below; veins distinct below; petiole 
stout or slender, 1-4 cm. Inflorescence up to 8 cm, 
slender or stout, subglabrous to densely appressed- 
hairy, up to 3(-4) times branched; branches of the 
first order scattered; bracts either all triangular, 
under 3 mm, or lower ones lanceolate, up to 15 
mm; basal inflorescence axes of the first order 
subtended by ordinary bracts or by leaves. 
Submature flower bud up to 3 1 2 mm Z. Sepals 4 or 
5, broader than long or as long as broad, thick, 
puberulous, to appressed-hairy. /Vm/s 4 or 5, thick, 
glabrous to appressed-hairy. Stamens 4-5. Ovary 
puberulous to appressed-hairy. Fruit ovoid to 
oblong, 2 1 4 -3' 2 by 1-1 l 4 cm; persistent disk con- 
spicuous or not; sepals inconspicuous. 

Distr. Continental SE. Asia (NE. India, 
Bhutan, Burma, Thailand, Tonkin, S. Yunnan) 
and throughout Malesia; not yet recorded from 
the Lesser Sunda Is. and New Guinea. 



KEY TO THE SUBSPECIES 

1. Flowers basically 4-merous. Inflorescence stout, 
rusty-puberulous. Corolla glabrous outside. 
Leaves obovate, coriaceous; apex acute; base 

attenuate b. ssp. moluccana 

1. Flowers basically 5-merous. 

2. Leaves up to 8-20 by 4-8 cm ; nerves 6 or more 

pairs; veins distinct beneath. Fruit ovoid. 

3. Basal bracts lanceolate, up to 15 mm. Fruit 

larger than 3 by l\ 4 cm . a. ssp. pentandra 

3. All bracts triangular, under 3 mm. Fruit up to 

2 l / 2 by 1 cm c. ssp. chinensis 

2. Leaves up to 4-12 by l\ 2 -5 cm; nerves up to 6 
pairs; veins obscure beneath. Fruit oblong. 



4. Length of sepals up to half as long as wide. 
Leaves chartaceous to subcoriaceous. Fruit 
2' 2 -3 cm long . . . . d. ssp. philippinensis 

4. Length of sepals almost as long as wide. 

Leaves coriaceous to thick-coriaceous. Bracts 

uniformly triangular. Fruit l 3 4 -2cm long 

e. ssp. scortechinii 

a. ssp. pentandra. 

Tree up to 34 m ; branchlets stout. Leaves elliptic 
to oblong-elliptic, 8-16 by 4-8 cm, coriaceous; 
base cuneate; apex acuminate; nerves 6-7(-9) 
pairs; veins distinct below; petiole stout, 2-4 cm. 
Inflorescence up to 8 cm, stout, densely appressed- 
hairy; basal bracts lanceolate, up to 15 mm. Sepals 
5, broader than long. Petals 5, appressed-hairy 
outside. Stamens 5. Fruit ovoid, 3-3 V 2 by IV4-IV2 
cm. 

Distr. Malesia: West & East Java. 

Ecol. In humid, mixed forest, 400-500 m. Fl. 
July-Dec. 

Vern. Java: huru HI in, tenjau. 

b. ssp. moluccana Matthew, Blumea 23 (1976) 81. 
Tree up to 15 m; d.b.h. 20 cm; branchlets stout. 

Leaves obovate, 8-15 by 3-8 cm, coriaceous; base 
attenuate; apex acute; nerves 5-6 pairs; veins 
distinct below; petiole stout, 2 1 2 -3 cm. Inflores- 
cence up to 5 cm, stout, rusty puberulous; basal 
bracts up to 4 mm. Sepals 4(-5), broader than long. 
Petals 4(-5), glabrous outside. Stamens 4(-5). 
Fruit (immature) ovoid, l 1 4 by 3 4 cm. 

Distr. Malesia: Moluccas (Morotai). 

Ecol. Mixed rain-forest, up to 1000 m. Fl. May. 
Once a leaf-gall was noted. 

Notes. The basal pair of lateral inflorescence 
branches often occur in the axils of normal leaves, 
a tendency noted in ssp. philippinensis. Flowers 
relatively large, yellowish to greenish; corolla 
dome-shaped (in bud). Calyx margin wavy; petals 
4 by 3 mm: filaments 3 mm; anther 1 mm; style 
1 ' 2 mm. The only fruit seen is detached and im- 
mature. 

The arrangement of the primary inflorescence 
branches is at times obscure. 

The soft, coriaceous texture of the leaves, dark 
above, and pale below, the stout inflorescence with 
rusty indumentum and the few, large, 4-merous 
flowers with glabrous dome-shaped corolla (in 
bud) distinguish this subspecies from the others. 
It is yet only known from Morotai I. 

c. ssp. chinensis (Merr.) Matthew, Blumea 23 
(1976) 83. — M. chinensis Merr. — Fig. 3c (galls). 

Tree up to 20 m; branchlets stout. Leaves 
elliptic to elliptic-oblanceolate, 8-20 by 4-8 cm, 
coriaceous; base attenuate; apex acute; nerves 6-8 
pairs; veins distinct below; petiole stout, l 3 / 4 - 
2 1 2 cm. Inflorescence up to 8 cm, subglabrous to 
appressed-hairy; all bracts uniform, under 3 mm. 
Sepals 5, broader than long. Petals 5, appressed- 
hairy outside. Stamens 5. Fruit oblong, 2-2V 2 by 
1 cm. 

Distr. NE. India, Bhutan, N. Burma, Thailand, 
S. China (Yunnan), Tonkin; in Malesia: Malay 
Peninsula (Kedah, once). 

Ecol. Mixed forests up to 1900 m. Fl. May- 
June, fr. Aug.-May. In India galls and domatia 
occur. 



1977] 



Cornaceae (Matthew) 



97 



d. ssp. philippinensis (Wangerin) Matthew, 
Blumea 23 ( 1 976)85. — M. philippinensis Wangerin 
— M. subcaudata Merr. — Fig. 3d-e (galls). 

Tree up to 22 1 2 m; brachlets slender. Leaves 
obovate to elliptic, 4-12 by l 1 2 -5 cm, chartaceous 
to subcoriaceous; base attenuate; apex acuminate; 
nerves 4-6 pairs; veins obscure beneath; petiole 
slender, 1-2 cm. Inflorescence up to 7 cm, slender, 
subglabrous to puberulous: all bracts triangular, 
under 3 mm. Sepals (4-)5, broader than long. 
Petals (4-)5, glabrous to appressed-hairy outside. 
Stamens (4-)5. Fruit oblong, 2 1 2 -3 by 1 ' 4 -l ' 2 cm. 

Distr. Malesia: throughout the Philippines. 

Ecol. In forests, from low altitude up to 1350 m. 
Ft. May-Sept., fir. Jan. -Dec. Galls are rather 
frequent on stem, leaf and fruit. 

Note. Ssp. philippinensis is distinguished from 
ssp. scortechinii in the generally smaller height and 
smaller and thinner leaves, more slender inflores- 
cence axis, the lower 1 or 2 inflorescence axis (axes) 
at the axil(s) of normal leaves, and the oblong 
fruits. 



e. ssp. scortechinii (King) Matthew, Blumea 23 
(1976) 86. — M. scortechinii King — M. mega- 
carpa Ridl. — M. parvifolia Hall. / 

Tree up to 37 m; branchlets stout. Leaves 
obovate to oblong, 4-12(-15) by 3-5(-6) cm, thick- 
coriaceous; base cuneate to attenuate: apex acute 
to acuminate; nerves 4-6 pairs; veins obscure 
beneath; petiole stout, l 1 2 -2' 2 cm. Inflorescence 
up to 8 cm, stout, puberulous to villous; all bracts 
triangular, under 3 mm. Sepals (4-)5, as long as 
broad. Petals (4-)5. Stamens (4-)5. Fruit oblong, 
2 1 4 -2' 2 by l J 4 -2cm. 

Distr. Thailand (once); in Malesia: S. Sumatra, 
Banka, Malay Peninsula, Borneo, Celebes. 

Ecol. Primary forests, from low altitude up to 



2400 m. Fl.fr. Jan.-Dec. Inflorences and fruit galls 
occur. 

Vern. Malaya: medang pisang. Banka: men- 
kapas. Borneo : kaju wulu, medang surugan. 

Note. Ssp. scortechinii is distinguished from 
ssp. pentandra by the generally obovate and smaller 
leaves, less stout inflorescence, uniformly short 
bracts and oblong fruits with thick pericarp. 

Excluded 

Mastixia cuneata Bl. Mus. Bot. Lugd. Bat. 1 
(1850) 257 = Notaphoebe umbelliflora (Bl.) Bl. 
Cf. Hall. f. Beih. Bot. Centralbl. 34, 2 (1916) 42; 
Danser, Blumea 1 (1934) 68. 

Mastixia gracilis King, J. As. Soc. Beng. 74, ii 
(1902) 73; Wangerin, Pfl. Reich Heft 41 4 (1910) 
28; Danser, Blumea 1 (1934) 68; Matthew, 
Blumea 23 (1976) 90 = Vaccinium bancanum Miq. 
var. tenuinervium J. J. S. {Ericaceae), according to 
the type number Wray 1528 mentioned by 
Sleumer, Blumea 11 (1961) 76. — Ed. 

Mastixia heterophylla Bl. Mus. Bot. Lugd. Bat. 
1 (1850)257; Wangerin, Pfl. Reich Heft 41 4 (1910) 
28; Danser, Blumea 1 (1934) 69; Matthew, 
Blumea 23 (1976) 90. — Hallier/. suggested this 
to be Gomphandra capitulata Becc, but this was 
questioned by Sleumer, Blumea 17 (1969) 193. 
According to us this sterile sheet (L 901, 169-350) 
collected by Praetorius in Palembang, is not a 
Mastixia but we cannot give a proper identification. 

Mastixia tetrandra (Thw.) Clarke. — Danser, 
Blumea 1 (1934) 56, referred two Sumatran speci- 
mens to this species, which is hitherto only found in 
Ceylon and the Andaman Is. One of these is 
sterile and the other is in bud ; they can equally well 
be referred to M. rostrata ssp. rostrata, and their 
identification remains doubtful. Cf. Matthew, 
Blumea 23 (1976) 77. 



Excluded 

Cornus caudata Hassk. & Zoll., nom. illeg., C. ilicifolia (Bl.) Hassk. & Zoll., C. serrulata (Bl.) 
Hassk. & Zoll., and C. stricta Zoll. & Mor. are all combinations or names made by Hasskarl and 
Zollinger based on Polyosma Bl., because of their opinion that this genus of the Saxifragaceae would be 
synonymous with Cornus L. Cf. Hasskarl, Cat. Hort. Bog. (1844) 168 and Zollinger, Natuur- & 
Geneesk. Arch. Neerl. Ind. 2 (1845) 10. 



ONAGRACEAE (P. H. Raven, St. Louis) 1 

Annual or perennial herbs (in Mai.), occasionally somewhat woody near the base, 
sometimes aquatic. Leaves spiral or opposite. Stipules absent or reduced, deltoid. 
Flowers mostly 4-merous, rarely 5-merous (in Mai.), solitary or arranged in a 
terminal racemose inflorescence, subtended by (often reduced) leaves or bracts. 
Bracteoles absent or 2 at the base of the ovary. Floral tube short or absent. Sepals 
erect, persistent. Petals caducous, contorted in aestivation, white, pink or yellow, 
sometimes emarginate. Stamens 4, 5, 8, or 10, in 2 whorls, rarely with an inter- 
mediate number, epipetalous ones sometimes shorter. Anthers usually versatile, 
sometimes seemingly basifixed by reduction: pollen single or in tetrads. Ovary 
inferior, (in Mai.) 4- or 5-celled and with cvi ovules; summit of the ovary (disk) flat 
to conical (in Mai.), sometimes with depressed nectaries surrounding the bases of 
the epipetalous stamens. Style simple; stigma capitate, clavate or globose, often 
4-lobed. Ovules with axial placentation, 1-pluriseriate. Fruit (in Mai.) a mostly long 
and slender loculicidal or irregularly rupturing capsule. Seeds rounded or elongate, 
in Ludwigia sometimes embedded in powdery or surrounded by cork-like endocarp 
tissue, in Epilobium with a chalazal plume of trichomes (coma); endosperm absent; 
embryo straight. 

Distribution. About 17 genera and more than 600 spp. in tropical and temperate regions, with a dis- 
tinct centre of diversity on the northern hemisphere in the New World, in Malesia two native genera which 
are both almost ubiquist. 

Ecology. Ludwigia is largely confined to the hot lowland and hills usually in wet or damp localities, 
Epilobium is confined to the higher mountain regions. 

Dispersal. Epilobium spp. are manifestly wind dispersed by virtue of their coma. Ludwigia spp. depend 
on dispersal by water and possibly incidental exozoic dispersal by water birds; in Ludwigia hyssopifolia 
there are two kinds of seed, one of which is enveloped by a corky tissue derived from the endocarp, 
enhancing their buoyancy. 

Pollination. Almost all of the Malesian species are self-pollinated, shedding pollen directly on the 
stigma at or before anthesis and rarely visited by insects. In Ludwigia peruviana, introduced in the Old 
World, the anthers are extrorse and shed pollen away from the stigma; thus outcrossing is predominant. 
Some outcrossing probably also occurs in the relatively large-flowered L. adscendens and L. octoyalvis, 
which are known to be visited by insects, and in Epilobium detznerianum, in some populations of which the 
stigma is even held above the anthers. In our area, Heide (Dansk Bot. Ark. 5, 1927, 18) reported Melipona 
sp., a bee, visiting the flowers of Ludwigia peruviana (as Jussieua peruviana), and Bombus rufipes at the 
flowers of the locally naturalized Oenothera stricta (as O. lamarckiana) and Fuchsia magellanica (as F. 
coccinea). The Melipona bees were not observed to contact the anthers or stigma of the large-flowered 
Ludwigia, but would certainly do so in visiting smaller-flowered species. All of the Malesian species are 
genetically self-compatible. 

Morphology & Anatomy. The Onagraceae are distinctive in their monosporic, 4-nucleate, 
"Oenothera type' embryo sac development: in the nearly universal presence of viscin threads among the 
pollen; and in the loose construction of their pollen exine. Most species of Epilobieae and about half of 
Ludwigia shed their mature pollen in tetrads; these include all Malesian species except L. adscendens and 
L. hyssopifolia, in which the pollen is shed singly. Intraxylary phloem occurs throughout the family 
adjacent to primary xylem, and interxylary phloem (included phloem) is found in many genera but not in 
Ludwigia (Carlquist, Ann. Mo. Bot. Gard. 62, 1975, 386); in these features Onagraceae resemble many 
other Myrtales. 

The stomata are surrounded by three or more subsidiary cells, sometimes resembling those of Cruciferae. 

All Onagraceae have an inferior ovary and a floral tube, which is prolonged beyond the ovary in all 
except Ludwigia and Epilobium sect. Chamaenerion. 

In several species of Ludwigia half-submerged parts of the stem are covered by a whitish aerenchyma; 
in L. adscendens short roots at the nodes are transformed into inflated, elongate aerophores enhancing 
floating on water. 

Chromosomes. All species of Epilobium sect. Epilobium, a taxon that includes all Malesian species, 
which have been examined have had a gametic chromosome number of n = 18. Species of Ludwigia have 
a gametic chromosome number of n = 8 and multiples. These genera differ from most others in Onagra- 

(1) With co-operation of the General Editor in framing the manuscript. 

The author gratefully acknowledges the support of the U.S. National Science Foundation to the studies 
of Onagraceae. 

(98) 



1977] Onagraceae (Raven) 99 

ceae in having small chromosomes that are heteropycnotic and dark-staining throughout the mitotic 
cycle. Naturally occurring interchange heterozygotes, abundant in the tribe Onagreae, are not known to 
occur in either group. The original basic chromosome number of the family is x = 11, as in Fuchsia, 
Circaea, and others. 

Hybridization. Hybrids are rare between the recognized species of Ludwigia. In Epilobium sect. 
Epilobium they are occasional where two or more entities come together, but their occurrence is limited by 
the predominant autogamy or cleistogamy of most species and to some extent by ecological differentiation 
also. A wide range of fertilities is characteristic of these hybrids, as explained in detail in our monograph 
of the Australasian species (Raven, D.S.I.R. New Zeal. Bull. 216, 1976), and cytoplasmic differences 
sometimes occur. Two of the Malesian species occur together in N. Luzon, and a few individuals suggest 
hybridization; four occur together in the mountains of New Guinea, with hybrids probably occasional 
but poorly studied so far. All species of Epilobium found in Malesia have the same chromosome arrange- 
ment that is predominant in Eurasia, from where they doubtless came. 

Chemotaxonomy. Raphides, needle-like crystals of calcium oxalate, are ubiquitous in the vegetative 
parts of Onagraceae. The few reports of alkaloids are doubtful and seem to indicate rather the presence of 
secondary amines. Ellagic acid occurs. Among the flavonoids reported from the family, flavonols based 
quercetin are ubiquitous, whereas kaempferol and more highly oxygenated types based on myricetin are 
frequent. The anthocyanins include predominantly malvidin and cyanidin derivatives, with the latter 
predominant in the rose-purple petals of Epilobium. The yellow petals of most species of Ludwigia are 
colored by carotenoids, with the chalcone isosalipurposide forming a non-ultraviolet-reflective centre in 
many species, including L. peruviana. 

Uses. Only some species of Ludwigia are mentioned to be in use for minor medicinal purposes; see 
under Ludwigia spp. 

KEY TO THE GENERA 

1. Capsule loculicidally dehiscent with 4 valves; axis persistent. Seeds comose. Floral tube present. 
Petals 4, white, pinkish or red. Stamens 8. Stem-base without aerenchyma 2. Epilobium 

1. Capsule irregularly dehiscent; axis not persistent. Seed not comose. Floral tube absent. Petals yellow, 
if white or creamy then flowers 5-merous and floating aquatic plant adorned with short spongy aeren- 
chyma-roots at the nodes. Stamens 4-10. Stem-base not rarely covered by aerenchyma. 1. Ludwigia 

1. LUDWIGIA 

Linne, Gen. PI. (1754) 55; Sp. PI. 1 (1753) 118; Munz, Bull. Torr. Bot. CI. 71 
( 1 944) 1 52 ; H ara, J. Jap. Bot. 28 ( 1 953) 289 ; A. & R. Fernandes, Garcia de Orta 5 
(1957) 109; Raven, Reinwardtia 6 (1963) 327. — Jussiaea Linne, Gen. PI. (1754) 
183; Sp. PI. 1 (1753) 388; Back. Trop. Natuur 3 (1914) 59; Fawcett, J. Bot. 64 
(1926) 10; Munz, Darwiniana4(1952) 179. — Nematopyxis Miq. Fl. Ind. Bat. 1, 1 
(1855)600. — Fig. 1,4, 5. 

Slender herbs, erect or creeping and rooting at the nodes, to large shrubs. 
Underwater parts often swollen and spongy or bearing inflated white spongy aero- 
phores. Leaves alternate or opposite, mostly entire. Stipules absent or reduced, 
deltoid. Flowers borne singly, clustered, or arranged in an inflorescence. Brac- 
teoles lacking or conspicuous, usually two, at or near the base of the ovary. Floral 
tube absent. Sepals 3-7, persistent after anthesis. Petals as many as the sepals or 
absent, caducous, yellow or white, with contorted aestivation. Stamens as many as 
or twice as many as the sepals, or flowers very rarely with an intermediate number 
of stamens; anthers usually versatile but sometimes apparently basifixed by reduc- 
tion. Pollen shed in tetrads or singly. Disk (summit of the ovary) flat to conical, 
often with depressed nectaries surrounding the bases of the epipetalous stamens. 
Stigma hemispherical or capitate, the upper l'/ 2 - 2 /., receptive, often lobed, the 
number of" lobes corresponding to the number of locules. Ovary with a number of 
cells equal to the number of sepals, very rarely more; placeiitation axial; ovules 
plunsenate or umsenate in each cell, in one species uniseriate below, pluriseriate 
above; if uniseriate, the seeds sometimes embedded in powdery or woody endocarp 



100 Flora Malesiana [ser. I, vol. 8 2 

from which they detach easily or with difficulty. Capsule irregularly dehiscent, or by 
a terminal pore, or by flaps separating from the valve-like top. Seeds rounded or 
elongate, the raphe usually easily visible and in some sections equal or nearly equal 
in size to the body of the seed. 

Distr. According to my synopsis (Reinwardtia 6, 1963, 329) 75 spp., all over the world; in Malesia 6 
spp., one of which is certainly introduced. 

Ecol. One aquatic and the other species mostly in swampy or damp places, often in rice-fields, from the 
lowland up to c. 2100 m, mostly below 1000 m. Flowers last only one day. 

Taxon. I have divided the genus into 17 sections, the largest of which (sect. Myrtocarpus) is neotropical. 
They are often shrubby with large, 4- or 5-merous flowers, dimerous stamens, prominently 4- or 5-ribbed 
capsules, free seeds and pollen grains shed in tetrads. They appear phylogenetically central in the genus. 
In Malesia this section is represented by an introduced weed, L. peruviana. Close to this section are one 
African (sect. Africana) and one American section (sect. Macrocarpori) with terete capsules. Following 
these is a series of small Old World sections which have the stamens reduced to one whorl ; in one African 
section flowers are 3-merous. L. hyssopifolia forms a monotypic section unique in having two kinds of 
seeds, those in the lower part of the capsule uniseriate and embedded in the endocarp, those in the upper 
part pluriseriate and free, while pollen grains are single, Other sections, not represented in Malesia, have 
all of the seeds loosely embedded in powdery endocarp. The structure of the seed is important in the dis- 
crimination of sections. 

The second major line of the genus consists of species in which the seeds are embedded in coherent 
chucks of woody endocarp which render the capsule a tough unit from which it is difficult to separate the 
seeds. The two sections belonging to this line have basically 5-merous flowers and pollen shed singly. 
Through the disentangling of these relationships it appears that the number of stamens is not decisive for 
dividing the genus into two genera as this would go across relationships and lead to heterogeneous assem- 
blages of species. 

Each Malesian species belongs to a different section and being so small in number it seems not useful to 
give descriptions of these sections; I refer to my revision (1963). 

The cradle of the genus is probably South America with an important secondary centre of evolution in 
Africa. It is one of the most primitive genera in the family. 

Note. It has appeared that seeds retain viability in the herbarium in unpoisoned, not too old speci- 
mens ; flowering plants can thus be raised from fruiting herbarium specimens. 



KEY TO THE SPECIES 

1. Stamens twice as many as sepals. 
2. Seeds pluriseriate, free (not embedded in endocarp). 
3. Plant subglabrous to appressed-pubescent. Capsule terete. Petals 5-17 by 4-17 mm. Style 1.5- 
3.5 mm. Raphe equal in diameter to the body of the seed 2. L. octovalvis 

3. Coarse, strongly pubescent or villous plant. Capsule strongly 4-angled, villous, with flat sides. 
Petals 15-24 by 16—26 mm. Style 1 mm long. Raphe not more than 1 U the diameter of the body of the 
seed 1. L. peruviana 

2. Seeds at least below uniseriate and embedded in endocarp. 

4. Seeds in the c. 1 U upper part of the capsule pluriseriate and free. Sepals 4. Petals 2-3 mm long 

5. L. hyssopifolia 
4. Seeds all uniseriate in each cell of the capsule and embedded in endocarp. Sepals 5-7 (rarely 4). Petals 
4.5-23 mm long. 
5. Aquatic, with floating branches forming erect clusters of spongy, spindle-shaped aerophores 

(aerenchyma). Petals white or creamy, with yellow at the base 6. L. adscendens 

5. Plant not forming such aerophores on the decumbent branches. Petals bright golden-yellow with a 
darker spot at the base. New Zealand, Australia, Pacific Is., Formosa, China, and Japan, also in the 
New World, might possibly be found in East Malesia. Cf. Aston, Aquat. PI. Austr. (1973) 144, f. 55 

L. peploides (H.B.K.) Raven 
1. Stamens as many as sepals, very rarely more in some flowers. 
6. Seeds pluriseriate in each cell of the capsule. Petals elliptical, 1-3 by 0.7-2 mm. Capsule terete. Seeds 

0.3-0.5 by 0.2-0.25 mm 3. L. perennis 

6. Seeds uniseriate in each cell of the capsule. Petals narrow spathulate, 1.3-2.2 by 0.4-0.9 mm. Capsule 
slightly 4-angled. Seeds 0.5-0.6 by 0.3 mm 4. L. prostrata 

1. Ludwigia peruviana (L.) Hara, J. Jap. Bot. 28 Steen. Fl. Sch. Indon. (1949) 305. — Oenothera 

(1953) 293; Raven, Reinwardtia 6 (1963) 345, hirta Linne, Sp. PI. ed. 2, 1 (1762) 491. — Jussieua 

map 14. — Jussiaea peruviana Linne, Sp. PI. 1 hirta (L.) Sw. Obs. Bot. (1791) 142, non Lamk, 

(1753) 388; Back. Trop. Natuur 3 (1914) 61 ; Onkr. 1789; Back. Ann. Jard. Bot. Btzg Suppl. 3 (1909) 

Suiker. (1930) 470, Atlas t. 445 ; Alston in Trimen, 406. — Jussiaea speciosa Ridl. J. Bot. 59 (1921) 

Fl. Ceyl.6(1931);Munz,Darwiniana4(1942)131; 259; Fl. Mai. Pen. 1 (1922)828. 



1977] 



Onagraceae (Raven) 



101 



Shrub 0.5-3 m, entirely villous, the hairs often 
multicellular, especially in the inflorescence; long 
inflated aerophores arising from submerged, 
buried roots. Leaves lanceolate to broadly lanceo- 
late, 4-12 by 0.3-1.5 cm, narrowly cuneate at base, 
apex acute to acuminate; nerves 12-22 pairs; sub- 
marginal vein not prominent; petiole 3-12 mm. 
Flowers in upper leaf axils. Bracteoles lacking or up 
to 7 mm long, subulate. Sepals 4 or 5, lanceolate, 
irregularly serrulate, 10-18 by 4-8 mm, villous. 
Petals bright yellow, veiny, suborbicular, 15-24 by 
16-26 mm, shallowly emarginate, with a claw 1-3 
mm. Stamens 8 or 10, subequal; filaments 2-3.5 
mm; anthers 3-4.5 mm long, extrorse and not 
shedding pollen directly on the stigma at anthesis. 
Pollen shed in tetrads. Disk elevated 1-2 mm, wi'h 
a depressed densely white-hairy nectary around the 
base of each epipetalous stamen. Style c. 1 mm; 
stigma broadly elongate-hemispherical, 2-3 mm 
high. Capsule villous, 1.2-3 by 0.6-1 cm, light 
yellowish brown with 4 prominent dark brown 
ribs, 4-angled, thin-walled, readily and irregularly 
loculicidal; pedicel 2-4.5 cm. Seeds pluriseriate in 
each cell, free, light brown, finely striate and cellu- 
lar pitted, obovoid, 0.6-0.8 mm long; raphe V4 to 
V 5 the width of the body. 

Gametic chromosome number (Old World 
populations), n = 40. 

Distr. Native of the New World, from the SE. 
United States throughout South America, intro- 
duced and naturalized in Malesia since the 2nd 
half of the last century, collected in Malaya, 
Sumatra, Java (common in West), but obviously 
still absent from many areas. 

Ecol. Along ditches and in moist places, mostly 
in the lowland but ascending to c. 1400 m. Fl. 
Jan. -Dec. 

Vern. Banka: pitjanket, M ; Java: tjatjabean, S, 
lombokan, J. 

Note. Backer (Onkr. Suiker. 1930, 470) 
observed that in inundated situation the plant 
produces aerophores which are emitted by shallow, 
horizontal roots; they are erect but their tips 
usually float on the water. 

2. Ludwigia octovalvis (Jacq.) Raven, Kew Bull. 
15 (1962) 476; Reinwardtia 6 (1963) 356, maps 
19-20, incl. ssp. brevisepala (Brenan) Raven et 
ssp. sessiliflora (Mich.) Raven; Henry & Prit- 
chard, Bot. Div. Lae, Bot. Bull. 7 (1973) 132, 
fig.; Everist, Pois. PI. Austr. (1974) 393. - 
Juulaea tuffruticosa I.inm', Sp. PI. 1 (1753) 388; 
Bin. Fl. Austr. 3 (1867) 307; F.v.M. Descr. Not. 
Pap. PI. 4 (1876) 60; Clarke, Fl. Br. Ind. 2 (1879) 
587; Koord. Exk. Fl. Java 2 (1912) 703; Ridl. 
Trans. Linn. Soc. Bot. II, 9 (1916) 57, incl. var, 
hirta Rim : GAON. Fl. Gen. I.-C. 2 (1921) 986; 
(I Whiii. Prut. R. Soc. Queensl. 34(1922)48; 
ft Baku. /. Fl, Java I (1963) 261. — Oeno- 
thera octoralvh Jacq. En. Syst. PI. (I7<>0) 19. - 

Sp. PI. ccl. 2, I (1762) 
555. — /.. perennU (/ion L.) Bi km. /. I I. Ind. (1768) 
37. — Jussiaea angustifolia Lamk, Encycl. 3 (I7K9) 
331. — Juulaea \ill<>\a Lamk, I.e.: Run . I I. Mai. 
Pen. I (1922) 82K. Juuleua octovalvii (J 
Sw Obs. Bot. (1791) 142. Juulaea angustifolia 
Bi . Bijdr. (1826) 1132, "<"> Lamk, 17X9 Baci 

I Natuur 3 (1914) 62. Jussiaea blunieana 

\x Prod. 3 (1828) 55. - Juulaea burmanntl DC. 



I.e. 57. — Jussiaea exaltata Roxb. (Hort. Beng. 
1814, 33, nomen) Fl. Ind. ed. Carey 2 (1832) 401. — 
Jussiaea costata Pr. Epim. Bot. (1849) 217. — 
Jussiaea junghuhniana Miq. Fl. Ind. Bat. 1, 1 (1855) 
627; Val. Bull. Dep. Agric. Ind. Neerl. 10 (1907) 
41. — Jussiaea erecta (non L.) Ridl. J. Bot. 59 
(1921) 258; Fl. Mai. Pen. 1 (1922) 827, incl. var. 
exaltata (Roxb.) Ridl.; Back. Onkr. Suiker. 
(1930) 470, Atlas t. 446; Steen. Arch. Hydrobiol. 
Suppl. 10 (1932) 314. — L. pubescens (L.) Hara, 
J. Jap. Bot. 28 (1953) 293. — Fig. 1. 

Usually robust, well-branched herb, sometimes 
woody at the base, up to 4 m, subglabrous or with 
sparse or dense appressed or spreading pubescence. 
Leaves lanceolate or narrowly lanceolate, to 
narrowly ovate, or subovate, 2-14 by 0.5-4 cm, 
narrowly to broadly cuneate at the base and 
attenuate at apex; nerves 11-20 pairs, sub- 
marginal vein well developed; petiole up to 1 cm. 
Bracteoles reduced or to 1 mm long. Sepals 4, ovate 
or lanceolate, 6-15 by 1-7.5 mm. Petals yellow, 
broadly obovate or cuneate, emarginate, 17 by 
2-17 mm. Stamens 8, epipetalous ones shortest; 
filaments 1-4 mm long; anthers 0.5-4 mm long, 
extrorse but soon crumbling and shedding pollen 
directly on the stigma. Pollen shed in tetrads. Disk 
slightly raised, with a white-hairy sunken nectary 
surrounding the base of each epipetalous stamen. 
Style 1.5-3.5 mm; stigma subglobose, shallowly 
4-lobed, 1.2-3 mm 0. Capsule thin-walled, 1.7- 
4.5 cm by 2-8 mm, terete, pale brown with 8 
darker ribs, readily and irregularly loculicidal; 
pedicel up to 10 mm. Seeds pluriseriate in each 
cell, free, brown, rounded, 0.6-0.75 mm long, 
0.5-0.7 mm wide including the inflated raphe 
which is equal in size to the body of the seed and 
evenly transversely ridged. 

Gametic chromosome numbers, n = 16, 24. 

Distr. Throughout the tropics of the world, 
between c. 32° N and 30 " S. 

Ecol. Mostly in humid places, damp grassland, 
rice-fields, along ditches and water-courses, in 
swamps, lakes and pools, drains, sandy or silty 
floodbanks, gravelly riverbeds, on floating islands 
in lakes, on floating logs in lagoons, sago swamps, 
mountain peat swamps with sedges, also in old 
native gardens and coconut plantings, from the 
lowland up to c. 1000 m, in Java and Celebes up to 
1400 m, in New Guinea up to 2100 m. Fl. Jan.- 
Dec. 

Several collectors mention that it propagates by 
runners and that old leaves turn reddish. The lower 
part of (he stem is at times coated by aerenchyma. 
In inundated condition aerophores are produced; 
sec under I . /.. peruviana. 

DOCTORS VAN LEEUWEN (Zoocecidia, 1926, 427 
f. X0X; Ned. Kruidk. Arch. 51, 1941, 204) recorded 
fruit galled by beetles and aphid galls on the ter- 
minal leaves of the branches. 

Uses. In Java minor medicinal qualities are 
ascribed to this species, amongst others against 

sprew. Rumphius, who described it under the name 
herba vttllgtnum (Herb. Amb. 6, p. 49) did not 

mention uses (Hivni, Nutt. PI. 1927. 1206). 

Burkili (Diet. 1935, 1274) reported that the 
mucilaginous leaves, aftei which the plant is called 
'liikom aye/ 'water 1 'ills', are used for poulticing 
in a variety ol complaints; it has also been recorded 
as used i"i headaches, orchitis, glands In the neck, 



102 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 1. Ludwigia octovahis (J acq.) Raven, a. Habit in flower, b. withered fruits, both x 1 I 2 (a after 

Backer, 1940, b Junghuhn ccxxii). 



1977] 



Onagraceae (Raven) 



103 



diarrhoea, dysentery, nervous diseases, and as a 
vermifuge. In Wilkinson's Dictionary it is said 
that a kind of tea is made from the leaves. Also in 
India medicinal properties are ascribed to it. 

Vern. Malaya: buyang samalam, lakom oyer, 
pujang malam, yenlidah, M ; Sumatra : urang aring 
it€m, Simalur I.; Java: gagabusan, tjatjabean, S, 
kalamenja, Md, salah njowo, J; Lesser Sunda Is.: 
pangambo, E. Sumbawa; Philippines: tayilakton, 
Tag., balansuit, Mag., malapdko, tubong-talapan, 
Bik., pachar-pachar, Sul., palangdisin, Ig., talang- 
kau. Ilk., halangot, naudyawa tubig, If.; Celebes: 
keletele tengen, Tonsaw. dial., Minahassa; 
Moluccas: daun panu, Ambon; New Guinea: ewo, 
onarenare, Kapauko lang., kampur, Sakaj bivouac, 
Merauke, pfauhanu, Kutubu, mayenke, Orne lang., 
Kaiye, rowijetwi, Enga lang., Yogos, togorarah, 
Wapi lang., Marok, rama-rama, Matapaili lang. 

Note. In my revision (1963) I distinguished three 
subspecies, more or less geographically defined, 
which I wish to withdraw here. 

3. Ludwigia perennis Linne, Sp. PI. 1 (1753) 119; 
A. & R. Fernandes, Garcia de Orta 5 (1957) 114, 
475; Raven, Reinwardtia 6 (1963) 367, map 21. — 
L. parviflora Roxb. (Hort. Beng. 1814, 11, nomeri) 
Fl. Ind. 1 (1820) 440; Bth. Fl. Austr. 3(1867)307; 
Kurz, J. As. Soc. Beng, 46, ii (1877) 91 ; Clarke, 
Fl. Br. Ind. 2 (1879) 588; Trimen, Fl. Ceyl. 2 
(1894)234; Ridl. Fl. Mai. Pen. 1 (1922) 828; Back. 
Onkr. Suiker. (1930) 471, Atlas t. 447; Back. & 
Bakh. /. Fl. Java 1 (1963) 261. — L. lythroides 
Bl. Bijdr. (1826) 1134. — L. gracilis Miq. Fl. Ind. 
Bat. 1, 1 (1855) 629. — Jussiaea perennis Brenan, 
Kew Bull. 8(1953) 163. 

Annual herb up to 1 m, subglabrous or minutely 
puberulent on younger parts. Leaves narrowly 
elliptic to lanceolate, 1-11 by 0.3-2.7 cm, narrowly 
cuneate at base, apex subacute; nerves 6-12 pairs; 
submarginal vein weakly developed; petiole 
2-15 mm, winged. Sepals 4, rarely 5, deltoid, 
(1.3-)2-3.5 by (0.5-)0.7-1.8 mm, glabrous or 
minutely puberulent. Petals yellow, elliptic, 1-3 by 
0.7-2 mm. Stamens usually 4 or 5, rarely more; 
filaments 0.3-0.7 mm; anthers 0.5-0.7 by 0.5- 
0.7 mm, shedding pollen directly on the stigma at 
anthesis. Pollen shed in tetrads. Disk slightly 
elevated, glabrous. Style 0.7-1.5 mm; stigma glo- 
bose, 0.4-0.5 mm 0. Capsule thin-walled, glabrous 
or puberulent, 3-16 (-19) mm long, terete, pale 
brown, readily and irregularly loculicidal; capsule 
sessile or on a pedicel up to 6 mm, often ! nodding. 
Seeds pluriseriate in each cell, free, brown with 
fine brown lines, ellipsoid-rounded, 0.3-0.5 by 
0.2-0.25 mm; raphe very narrow and inconspi- 
cuous. 

Gametic chromosome number, n = 8. 

Distr. Tropical Africa (from Senegal, Lake 
Chad, and the Sudan south to E. Congo and N. 
Natal), Madagascar, through continental SE. Asia 
(Ceylon to S. ( hina, Hongkong, and Hainan), and 
throughout Malesia (except Borneoand Moluccas) 
to tropical Australia and New Caledonia. Fig. 2. 

! Sunny, humid or marshy situations 
fallow rite-fields, sawah dikes, along ditches, rivers 
and water-courses, dry riverbeds, sugarcane fields 
on heavy clay, damp plates in I UCalypI savannah 
(New Guinea), from the lowland l<> c. 900m, II 
Jan. Dec. 



In Java and Madura I. decidedly preferring 
regions subject to a dry monsoon. 




Fig. 2. Range of Ludwigia perennis L. (after Raven, 
1963). 

4. Ludwigia prostrata Roxb. (Hort. Beng. 1814, 1 1, 
nomeri) Fl. Ind. 1 (1820) 441; Bth. Fl. Austr. 3 
(1866) 308; Clarke, Fl. Br. Ind. 2 (1879) 588; 
Trimen, Fl. Ceyl. 2 (1894) 234; Merr. Fl. Manila 
(1912) 355; Ridl. Fl. Mai. Pen. 1 (1922) 829; 
Merr. En. Philip. 3 (1923) 220; Back. Onkr. 
Suiker. (1930) 472, Atlas t. 448; Back. & Bakh. 
/. Fl. Java 1 (1963) 261; Raven, Reinwardtia 6 
(1963) 374, map 23. — L. fruticulosa Bl. Bijdr. 
(1826) 1133. — L. leucorhiza Bl. I.e. — Nema- 
topyxis pusitla Miq. Fl. Ind. Bat. 1, 1 (1855) 630. — 
Nematopyxis prostrata Miq. I.e. — Nematopyxis 
fruticulosa MlQ. I.e. 

Annual herb 0.1-0.6 m, subglabrous, often 
reddish-tinged. Leaves elliptic or narrowly elliptic, 
1-13 by 0.3-2.7 cm, glabrous or with a few minute 
hairs along the veins, narrowly cuneate at the base, 
apex acute; submarginal vein weakly developed; 
petioles 4-25 mm, distinct. Sepals 4, deltoid, 1.3- 
2.5 by 0.7-1 . 1 mm, glabrous. Petals yellow, narrow- 
ly spatulate, 1.3-2.2 by 0.4-0.9 mm. Stamens 4; 
filaments 0.8-1.2 mm; anthers 0.4-0.5 mm wide, 
broader than long, closely appressed to the stigma 
and shedding pollen directly on it at anthesis. 
Pollen shed in tetrads. Disk slightly elevated, 
glabrous. Style c. 1 mm ; stigma globose, c. 0.5 mm, 
the upper half receptive. Capsule thin-walled, 
glabrous, 12-22 by 0.8-1 mm, { 4-angled, pale 
brown, readily and irregularly loculicidal, the 
seeds showing plainly as indentations in the walls 
at maturity. Seeds uniseriate in each cell, free, pale 
brown, speckled or striped transversely with narrow 
darker brown stripes, plump, ovoid, apiculate at 
one end, 0.5-0.6 by 0.3 mm; raphe narrow, linear. 

Gametic chromosome number, n = 8. 

Distr. Tropical SE. Asia (Ceylon and S. Dcccan 
to N. India, Assam, S. China and Andamans); in 
Malesia: Malay Peninsula, Java, Lesser Sunda Is. 
(Timor), Hornet), and Philippines (Palawan, 
Luzon, Negros, Mindanao). The single record from 
Australia (N. Queensland: Mossman R.) recorded 
I . Win 1 1 (Proc. R. Soc. Queensl. 50, 1939, 
78) is a misulentilication of / . hysSOptfblia. On the 
whole /.. pmtrata is in Malesia a rare species. 
Fig. 3. 

I col. Paddies, fallow and planted, by ditches. 
ftlonfl rivcrhanks, in svvampv places, rather rare. 

from lowland up to c. BOO m. //. Jan. Oct, 

Vi kn. Philippines: alubihud, P. His. 



104 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 3. Range of Ludwigia prostrata Roxb. (after 
Raven, 1963). 

5. Ludwigia hyssopifolia (G.Don) Exell, Garcia 
de Orta 5 (1957) 471 ; A. & R. Fernandes, I.e. 471, 
474; Raven, Reinwardtia 6 (1963) 385, map 30; 
Henty & Pritchard, Bot. Div. Lae, Bot. Bull. 
7 (1973) 131, fig. — Jussiaea linifolia Vahl, Eclog. 
Am. 2 (1798) 32, non L. linifolia Poir. 1813; Back. 
Trop. Natuur 2 (1913) 20, fig.; ibid. 3 (1914) 61; 
Onkr. Suiker. (1930) 469, Atlas t. 444; Steen. 
Arch. Hydrobiol. Suppl. 10 (1932) 314; Back. & 
Bakh./. Fl. Java 1 (1963) 260. — Jussiaea hyssopi- 
folia G.Don, Gen. Syst. 2 (1832) 693. — Jussiaea 
suffruticosa {non L.) Ridl. J. Bot. 59 (1921) 257; 
Fl. Mai. Pen. 1 (1922) 827. — L. prostrata {non 
Roxb.) C. T. White, Proc. R. Soc. Queensl. 50 
(1939) 78. — Fig. 4. 

Annual herb 5 cm to 3 m, often becoming woody 
at the base; young growth and inflorescence 
minutely puberulent; elongate aerophores arising 
from buried submerged roots. Leaves lanceolate, 
1-9 by 0.2-3 cm, narrowly cuneate at the base, 
apex acuminate; nerves 11-17 pairs; submarginal 
vein not prominent; petiole 2.5-18 mm. Sepals 4, 
lanceolate, 2-4 by 0.7-1.2 mm, finely puberulent, 
3-nerved. Petals yellow, fading orange-yellow, 
elliptic, 2-3 by 1-2 mm. Stamens 8, pale greenish 
yellow, epipetalous ones shorter; filaments of 
episepalous stamens 1-2 mm, those of epipetalous 
ones 0.5-1 mm; anthers 0.4-0.6 mm wide, 0.2- 
0.3 mm high, shedding pollen directly onto the 
stigma at anthesis. Pollen grains shed singly. Disk 
± elevated, with a depressed ciliate nectary sur- 
rounding the base of each epipetalous stamen. 
Style pale greenish yellow, 1-1.5 mm; stigma de- 
pressed-globose, c. 0.6-1.2 mm 0, 0.5-0.8 mm 
high, shallowly 4-lobed, the upper portion recep- 
tive. Capsule relatively thin-walled, finely puberu- 
lent, 1.5-3 cm by 1-1.2 mm, subterete, enlarged 
in the upper J 6 to 1 3 , subsessile. Lower seeds uni- 
seriate in each cell of the capsule, nearly vertical, 
brown, oblong, 0.7-0.85 mm long, each firmly 
embedded in a cube of relatively hard endocarp; 
raphe ' 3 the diameter of the body. Seeds in upper 
inflated portion of the capsule multiseriate, free, 
ovoid, 0.35-0.5 mm long, paler brown than the 
lower seeds and with a narrower raphe. Lower part 
of capsule at first marked by distinct bumps 
corresponding to the position of the uniseriate 
seeds, but as the endocarp hardens and swells, the 
capsule becomes smooth. 



Gametic chromosome number, n = 8. 

Distr. Tropical Africa (Dakar to Lake Chad, 
S. Sudan and S. Congo), continental SE. Asia 
(Ceylon to Hainan), throughout Malesia to 
Micronesia and N. Australia (Cape York Penin- 
sula and Arnhem Land). 

It is difficult to fix the native country of this now 
widely spread palaeotropical weed which has no 
close relatives. It might have been introduced in 
tropical Africa where it is relatively local and con- 
fined to the west, but it was collected in Sao Tome 
as early as 1822. 

Ecol. A very common weed of pools, along 
drains, shallow ditches, water and river edges, in 
paddies and humid, fallow garden land, on waste 
land, fallow sawahs, and in clearings, both on clay 
and humid white sand (Borneo), in pools in Euca- 
lypt savannah (Wetar I.), on Mt Kelud (Central 
Java) as a pioneer on old volcanic mudstreams 
(lahars), from the plains up to c. 1000 m. Fl. Jan- 
Dec. 

Backer (Trop. Natuur 2, 1913, 133) described in 
detail the biology of this species, of which the 
flowers open at 7 a.m. Seeds are gradually released 
by decay of the pericarp of which the vessels remain 
persistent. To his surprise both kinds of seed were 
buoyant for 16 days, after which they sank. On the 
17th day he observed one buoyant seedling, but 
unfortunately no mention was made from which 
kind of seed; his experiment was through inter- 
ference discontinued. He made also notes on the 
aerophores emitted by the shallow roots as occur 
also in L. peruviana and other species. 

Docters van Leeuwen (Zoocecidia, 1926, 428; 
Ned. Kruidk. Arch. 51, 1941, 204) reported galled 
fruits caused by beetles. 

Leaves are often attacked by a blue beetle: 
Craptodera cvanea (Backer, Trop. Natuur 2, 1913, 
132). 

Uses. Heyne (Nutt. PI. 1927, 1206) noted that 
in N. Celebes it is used for poulticing pimples. 
According to Burkill (Diet. 1935, 1273) it is in 
Malaya generally stocked by Chinese herbalists, 
but its use is not clear; it was once recorded that 
an infusion of the root is swallowed by Malays for 
syphilis. In the Philippines the whole plant is used 
for black dye (Quisumbing, Medic. PI. Philip. 
1951,676). 

Vern. Sumatra: meligai, M, Banka: Java: 
(djukut) anggereman, mainang, tjatjabean, S; 
Philippines: pasau-na-hapai, sila sila, Tag., dam- 
num- wiliyan, Mag., kakaggin diloba, If., barigaud, 
Bik., manakatud, Ilk., talang-duron, Pamp., 
tohod-tohod, Bik. ; Celebes: kayu ragi, Manado. 

Note. The dimorphous seeds are very unusual 
and it would be most interesting to have informa- 
tion on the properties of the two seed types with 
respect to germination. 

6. Ludwigia adscendens (L.) Hara, J. Jap. Bot. 28 
(1953) 290; A. & R. Fernandes, Garcia de Orta 5 
(1957) 475; Raven, Reinwardtia 6 (1963) 387, maps 
31, 33; Aston, Aquat. PI. Austr.(1973) 142; Henty 
& Pritchard, Bot. Div. Lae, Bot. Bull. 7 (1973) 
130, fig. — Jussiaea repens Linne, Sp. PI. 1 (1753) 
388, nonL. repens Forst. 1771 ; F.v.M. Descr. Not. 
Pap. PI. 4 (1876) 60; Clarke, Fl. Br. Ind. 2 (1879) 
587; O. K. Rev. Gen. PI. 1 (1891) 251, inch var. 
pilosa O. K. et var. glaberrima O. K. ; Trimen, Fl. 



1977] 



Onagraceae (Raven) 



105 




Fig. 4. Ludwigta hystopifolia (Q Don) i keli i. Habit, in flower, b. withered fruits, both ' i(aaftei 

H\. ker, 1940, 6 Verboom 9). 



106 



Flora Malesiana 



[ser. I, vol. 8 2 



Ceyl. 2 (1894) 233; Koord. Exk. Fl. Java 2 (1912) 
703; Back. Trop. Natuur 3 (1914) 56, 60, f. 1-5; 
Ridl. Fl. Mai. Pen. 1 (1922) 827; Gagn. Fl. Gen. 
I.-C. 2 (1925) 987 ; Hochr. Candollea 3 (1925) 479, 
incl. ssp. glabrata Hassl. /. albiflora Hochr.; 
Hartsema, Flora (Allg. Bot. Z). n.s. 22 (1927) 242, 
t. 3; Back. Onkr. Suiker. (1930) 469, Atlas, t. 443; 
Steen. Arch. Hydrobiol. Suppl. 10 (1932) 314, 
f. 62; Fl. Sch. Indon. (1949) 305; Back. & Bakh. 
f. Fl. Java 1 (1963) 260. — Jussiaea adscendens 
Linne, Mantissa 1 (1767) 69. — Jussiaea fluviatilis 
Bl. Bijdr. (1826) 1132. — Fig. 5. 

Herb with prostrate or ascending stems, rooting 
at the nodes, with conspicuous white, erect, 
spindle-shaped, mucronate aerophores arising in 
clusters at the nodes of the floating stems and from 
the roots, the more or less erect stems to 60 cm ; 
floating stems to 4 m; plants normally glabrous, 
but the branches growing on dry ground densely 



villous and rarely flowering. Leaves broadly 
oblong-elliptical, 0.4-7 by 0.7-4 cm, narrowly 
cuneate at base, apex acute or obtuse; nerves 6-13 
pairs; submarginal vein not prominent; petioles 
long. Flowers in upper leaf axils. Bracteoles 
present near base of capsule, deltoid, c. 1.2 by 1.3- 
1.5 mm. Sepals 5, deltoid-acuminate, 5-11 by 
2-3.2 mm, glabrous or villous. Petals creamy white, 
yellow at base, obovate, rounded at apex, 9-18 by 
6-10 mm. Stamens 10, epipetalous ones slightly 
shorter; filaments white, 2.5-4 mm; anthers 
1.2-1.8 mm long, apparently shedding pollen 
directly on the stigma at anthesis. Pollen grains 
shed singly. Disk ± elevated, with a depressed 
white-hairy nectary surrounding the base of each 
epipetalous stamen. Style white, 4-8 mm, densely 
long-hairy in lower half; stigma globose, green, 
1.5-2 mm 0, 1-1.2 mm high, the upper 2 / 3 recep- 
tive. Capsule glabrous or villous, 1.2-2.7 cm by 




Fig. 5. Ludwigia adscendens (L.) Hara, x 2 / 3 (after Backer, 1940). 



1977] 



Onagraceae (Raven) 



107 



3-4 ram, light brown, with 10 conspicuous darker 
brown ribs, terete, the seeds evident between the 
ribs as bumps c. 1.5 mm apart; capsule thick- 
walled, very tardily and irregularly dehiscent; 
pedicel 2.5-5.5 cm. Seeds uniseriate in each cell of 
the capsule, pale brown, 1.1-1.3 mm long, ± 
vertical, firmly embedded in coherent cubes of 
woody endocarp 1 .2-1.5 mm high, 1-1.2 mm thick, 
the endocarp firmly fused to the capsule wall. 

Gametic chromosome numbers, n = 8, 16. 

Distr. Continental Asia (from Ceylon to S. 
China), throughout Malesia, in Australia one 
locality in W. Arnhem Land. Fig. 6. 

Ecol. Freshwater pools, ditches, swamps, fallow 
and wet paddies, very common, from the lowland 
up to c. 1600 m. Fl. Jan.-Dec. 




Fig. 



6. Range of Ludwigia adscendens (L.) Hara 
(after Raven, 1963). 



Backer (Trop. Natuur 3, 1914, 56) depicted and 
described in detail the biology. The root system 
consists of three kinds, long ± unbranched anchor 
roots, shorter much-branched feeding roots, and 
the erect spongy aerophores. Backer cut the latter, 
but the plant remained (only very slightly less) 
buoyant. After pollination the pedicel bends down 
and the fruit ripens in the water (as in several other 
aquatics); the fruit decays and releases the cork- 
winged seeds which are buoyant. 

On desiccated muddy soils a never-flowering 
terrestrial form often occurs, marked by very small 
crowded leaves and a stronger pubescence. 



Uses. Malays in Perak use it for poulticing in skin 
complaints (Burkill, Diet. 1935, 1273). Batak 
people use this (and other aquatics, like also do the 
Chinese) to feed pigs, and it is recorded to be eaten 
as salad in Indo-China. Quisumbing (Medic. PI. 
Philip. 1951, 677) reported that it is used in a 
decoction as an astringent for dysentery. 

Vern. Water primrose, E; Sumatra: buang 
buang, Toba-Batak; Java: pangeor, M, gangging 
landeuj, kambang peutjit, ruba silah, (rumput) 
kologa, S, krangkong, krema, patjar banju, tapak 
doro, J; Philippines: sigang-ddgat, Tag., gdbi-gabi, 
Mag., tabagan, If.; New Guinea: agidahano, 
Kutubu lang. 

Note. L. adscendens appears to be allied more 
closely to the American L. helminthorrhiza (Mart.) 
Hara than to any Old World species. Together 
with the mostly yellow-flowered African L. 
stolonifera (Guill. & Perr.) Raven, these three 
are the only species that regularly produce clusters 
of erect inflated aerophores at the floating nodes, 
although other species have descending root-like 
aerophores at these nodes and may have long, 
spongy aerophores from the submerged under- 
ground parts. 



Excluded or Doubtful 

Jussiaea tenella Burm./. Fl. Ind. (1768) 103, t. 34, 
f. 2. 

There is no unanimity of opinion about the 
identity of Burman's plant which he said to have 
come from Java. Merrill (Philip. J. Sc. 19, 1921, 
369) suggested it to be a form of L. octovalvis which 
I doubt in view of the specific epithet and the 
species with which it is being contrasted. Alston (in 
Trimen, Fl. Ceyl. 6, 1931, 130) took it up for 
L. hyssopifolia, and this suggestion seems the most 
plausible. From Burman's description, which 
contains almost certainly errors, it cannot be 
identified. Unfortunately, or perhaps fortunately, 
the type at G could not be found. 

Ludwigia erigata Linne, Mantissa 1 (1767)40. — 
L. triflora Desr. in Lamk, Encycl. 3 (1792) 615, 
nom. illeg. subs., belongs to the Rubiaceae. 

Ludwigia trifolia Burm. /. Fl. Ind. (1768) 36; 
Houtt. Nat. Hist. 2, 7 (1777) 344 is according to 
Merrill (J. Am. Arb. 19, 1939, 368) Oldenlandia 
biflora L. (Rubiaceae). 



2. EPILOBIUM 

Linm. Gen. PI. ed. 5 (1754) 164; Sp. PI. 1 (1753) 347; Hausskni;cht, Monogr. 
Epilob. (1884); Ravin, Bull. Br. Mus. Nat. Hist. Bot. 2 (1962) 325, 13 maps, pi. 33- 
39; BKorKiF., New Zeal. J. Bot. 4 (1966) 366, 2 fig.; Bothalia 9 (1967) 309, 7 fig.; 
Ravf.n, Blumea 15 (1967) 269, 7 fig.; Bko< kii. New Zeal. J. Bot. 8 (1970) 94; 
Ravin, D.S.I. R. New Zeal. Bull. 216 (1976) 321 pp., 158 fig. Fig. 7. 

Perennial herbs, often flowering in the first year, occasionally somewhat woody 
near the base. Leaves (in Mai.) opposite below, spirally arranged above, flowers in 
the axils of the often greatly reduced upper leaves, floral tube short (in Mai.) or 
essentially absent. Sepals 4, erect. Petals 4, while, pink, or purple, emarginate 



108 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 7. Epilobium hooglandii Raven, a. Habit, nat. size, b. node, x 10, c. seed with base of coma, x 30. — 
E. detznerianum Schltr ex Diels. d. Node, x 10. — E. prostratum Warb. e. Seed, x 30 (a-c Hoogland 
& Pullen 5540, d Hoogland & Pullen 5687, e Womersley c.s. 6103). After Raven, 1967. 



1977] Onagraceae (Raven) 109 

(in Mai.)- Stamens 8, in 2 whorls, the epipetalous ones shorter. Stigma (in Mai.) 
clavate or rarely globose, usually surrounded by the shedding anthers at maturity. 
Ovary 4-locular, the ovules very numerous. Fruit a long, slender, loculicidal cap- 
sule. Seeds very numerous, small, with a chalazal plume of trichomes (coma). 

Distr. About 200 spp., well-represented in temperate regions, mostly on the northern hemisphere, with 
the greatest centre of morphological diversity in the western U.S.A., at relatively high latitudes and alti- 
tudes; in the tropics confined to the mountains; in Malesia: rare and local, in W. Central Sumatra, E. Java 
(Mt Tengger), Lesser Sunda Is. (Lombok, Sumbawa, Timor), Philippines (N. Luzon, Panay), SW. 
Central Celebes (Latimodjong Range), Moluccas (Central Ceram), and New Guinea. 

Ecol. Characteristic of open, disturbed habitats or grassland or the alpine zone, not normally found in 
primary forest. 

Dispersal takes easily place by wind as the light seeds are provided with a silky coma. 

It is still rather surprising that, whereas open habitats are very common in the Malesian mountains, 
with their numerous volcanoes and alpine habitats, the number of localities is so very restricted. It must 
probably be considered that though dispersal may be easy, the coma only acts efficiently in dry air and this 
is in the tropics with their frequent rain and cloud formation and nightly fogs during most of the year a 
rather rare climatic situation. In this respect the west-monsoon, blowing from Asia southeastwards (Nov.- 
March) is most unsuitable, as these winds are laden with moisture. The east-monsoon, blowing from 
Australia northeastwards (May-Sept.) is in this respect definitely more favourable as this is a dry wind. 
This wind regime is a consequence of the situation of the Asian and Australian continents. 

Local dispersal by water might also occur, as the lowest localities in New Guinea are all along stream- 
sides, but this may also be due to the downward air-stream over rushing water. 

For hybridization see under the family. 

Genesis. The species occurring in Sumatra (1 sp.) and in Luzon (2 spp.) also occur in continental SE. 
Asia, the one in E. Java and the Lesser Sunda Is. (E. hirtigerum) also occurs in Australia, Tasmania, and 
New Zealand. The four remaining species are all endemic in East Malesia, 3 being confined to New Guinea 
and 1 also occurring in Central Celebes and Ceram. They are closely related to species found in Australia 
and New Zealand, from which they, and E. hirtigerum, were probably derived via the east-monsoon dis- 
cussed above. 

I have argued that the Australasian species ultimately show affinity to those of continental Asia and 
their ancestors must have reached Australia across the tropical mountains of Malesia. My assumption is 
that this southeastward penetration of the genus occurred in the Pliocene. 

KEY TO THE SPECIES 

1. Buds and flowers erect, the inflorescence erect or somewhat drooping. 
2. Stems pubescent only along conspicuously elevated lines running down from the margins of the 

petioles 3. E. wallichianum 

2. Stems pubescent all around. 
3. Upper leaves strongly reduced. Inflorescence slightly nodding (Luzon). 
4. Leaves ovate, 0.4-1.5 cm wide 1. E. brevifolium 

4. Leaves very narrowly elliptic to linear, 0.1-0.5 cm wide 2. E. platystigmatosum 

3. Upper leaves not notably reduced. Inflorescence erect (Java, Lesser Sunda Is., New Guinea). 

5. Plants finely glandular-pubescent; leaves usually ericoid in appearance; petals purplish-rose 

5. E. keysseri 
5. Plants densely strigulose; leaves broader, not ericoid; petals white or very pale pink 

4. E. hirtigerum 
I. Buds and flowers drooping, becoming erect in fruit. 
6. Stems with elevated glabrous lines running from the center of each petiole and elevated pubescent lines 

from their angles; petals 7-14 mm long 6. E. detznerianum 

6. Stems without elevated lines, uniformly pubescent all around; petals 2.5-8.5 mm long. 

7. Petals 6-8.5 mm long; seeds 1-1.4 mm long 7. E. hooglandii 

7. Petals 2.5 5( -6) mm long; seeds 0.7-0.9 mm long 8. E. prostratum 

I. Kpilobium brevifolium I). Don, Prod. Fl. Nepal. Robust, perennial herb, !5-40cm, the under- 

(1825) 222; Raven, BuU. Br. Mus. Nat. Hist. Bot. 2 ground parts not scaly; plants strigulose, stems 
(1962)361. pubescent all around. Leaves mostly opposite, 

alternate in and near the inflorescence, the upper 
\\p. trichoneurum III- KNECHT) Ravkn, Bull. ones reduced, strigulose along the veins and 
Mr. Mus. Nat. Hist. Bot. 2 (1962) J62. / tricho- margins especially below, the nerves evident, ovate, 

newrum Hmsskmmii, Oest. Bot. Z. 29 (1879) 54; I 3 by 0.4 l.5cm, serrate; petiole I 2 mm. In- 

.gr I pilob. (1884)208; H. LivEiLii, Ic. Gen. florescence slightly nodding. / lowers erect, borne 

l pilob. (1910) t 84. / phtllpplnense C.B. R<>» in the axils of reduced upper leaves, floral tube 

Philip. J. Sc. 3 (1908) Bot. 209; Mere. En. Philip. 3 1.8 2.4 mm across, i 1.3 mm deep. Sepals 2.3 5.5 

M'>2i) 220. exel. tyn, by l 1.3 mm. Petals obovate, 4.5 8 by 3 5.5 mm, 



110 



Flora Malesiana 



[ser. I, vol. & 



rose purple, the notch 1-1.5 mm deep. Anthers 
1-1.2 mm long; filaments of the longer stamens 
1.2-2.5 mm, those of the shorter 1-1.5 mm. Style 
2.5-4 mm. Stigma clavate, 1.8-2.3 mm high, c. 
1 mm thick, surrounded by the anthers at anthesis. 
Capsule 3.5-7 cm long, on a pedicel 0-1 cm. 
Seeds 0.9-1.2 by 0.4-0.5 mm, papillose, blackish 
brown, not beaked, obovoid, blunt at both ends, 
the coma 5-7 mm long, white. 

Gametic chromosome number, n = 18. 

Distr. Continental SE. Asia (SE. Tibet: 
Chumbi Valley, to Assam, Burma, and W. China); 
in Malesia: Philippines (Mountain Province of 
N. Luzon). 

Ecol. Wet open slopes in the pine region, along 
streams and by springs, 1400-2100 m. Fl. July, Oct. 

Note. E. brevifolium ssp. trichoneurum is one of 
three subspecies of a species which ranges from 
Himachal Pradesh in the Western Himalaya east- 
wards throughout the Himalaya and southern 
China to Formosa, northern Luzon, northern 
Vietnam, and northern Burma. In northern Luzon, 
it occurs together with the other species found in 
the Philippines, E. platystigmatosum, and one plant 
of the collection Clemens 16385, suggests hybridi- 
zation between these two entities, which are usually 
widely distinct morphologically, although doubt- 
less more closely related to one another than to 
other Malesian species. 

2. Epilobium platystigmatosum C. B. Robinson, 
Philip. J. Sc. 3 (1908) Bot. 210; Merr. En. Philip. 3 
(1923) 221; Raven, Blumea 15 (1967) 272. — 
E. cephalostigma var. linearifolium Hisauti, J. Jap. 
Bot. 14(1938) 143, f. 3. — E.formosanum Masum. 
Trans. Nat. Hist. Soc. Formosa 29 (1939) 62; 
Ohwi, Fl. Japan (1965) 657. — E. sohayakiense 
Koidz. Act. Phytotax. Geobot. 8 (1939) 61. 

Slender, perennial herb, 1 5-40 cm tall, the under- 
ground parts not scaly ; plants strigulose, densely so 
in the inflorescence, the stems pubescent all around. 
Leaves mostly opposite, alternate in and near the 
inflorescence, the upper ones reduced, strigulose 
along the veins and margins especially below, the 
nerves evident, very narrowly elliptic or linear, 
1— 3(-4) by 0.1-0.5 cm, weakly and sparsely ser- 
rulate; petiole l-4mm. Inflorescence slightly 
nodding. Flowers erect, borne in the axils of reduced 
upper leaves. Floral tube c. 1 mm across, c. 0.8 mm 
deep. Sepals 3-4 by 1.2-1.6 mm. Petals narrowly 
obovate, 3.3-4.5 by c. 2 mm, white or pale pink, 
the notch c. 1 mm deep. Anthers 0.2-0.3 mm long; 
filaments of the longer stamens c. 1.8 mm, those of 
the shorter c. 1 .2 mm. Style c. 2 mm. Stigma 
broadly clavate, c . 0.8 mm high, c. 0.4 mm thick, 
surrounded by the anthers at anthesis. Capsule 
glabrescent, 3-5 cm long, on a pedicel 0-1.8 cm. 
Seeds 0.8-1 by 0.3-0.4 mm, papillose, not beaked, 
obovoid, blunt at both ends, the coma 4-6 mm 
long, white. 

Gametic chromosome number, n = 18. 

Distr. Japan, China (Hupeh, Kiangsu), For- 
mosa, and Malesia: Philippines (N. Luzon: 
Benguet Prov.; Panay: BS 31439). 

Ecol. Chiefly along small streams and about 
cliffs, 1200-2400 m. Fl. April-June. 

Note. I can find no difference between the species 
generally known as E. formosanum and the sup- 
posed Philippine endemic populations of E. philip- 



pinense. This species is apparently not common on 
the Asian mainland. 

3. Epilobium wallichianum Haussknecht, Oest. 
Bot. Z. 29 (1879) 54; Raven, Bull. Br. Mus. Nat. 
Hist. Bot. 2 (1962) 365; Blumea 15 (1967) 272. — 
E. nepalense Haussknecht, Oest. Bot. Z. 29 (1879) 
53, p.p.; Monogr. Epilob. (1884) 218, p.p.; H. 
Leveille, Ic. Gen. Epilob. (1910) t. 120. — 
E. duclouxii H. Leveille in Fedde, Rep. 6 (1908) 
110; Ic. Gen. Epilob. (1910) t. 144. — E. sarmen- 
taceum (non Haussknecht) Bunnemeuer, Trop. 
Natuur 10 (1921) 57, f. 9. 

Erect perennial herb 1 5-50 cm, from a long 
rhizomatous base from which leafy shoots arise; 
plants strigulose, more densely so above, with 
elevated pubescent lines running down from the 
sides of the petioles, stems thick and hollow. Leaves 
opposite in lower half of the plant, alternate 
above, the margin and nerves densely strigulose, 
narrowly ovate or lanceolate, subacute at the apex, 
obtuse at the base, sharply and densely serrulate, 
1.5-4 by 0.5-1.5 cm; petiole short but distinct, up 
to 2 mm. Inflorescence densely strigulose with an 
admixture of glandular trichomes, somewhat 
nodding in bud. Floral tube c. 2 mm across, c. 
0.8 mm deep. Sepals c. 5 by c. 2 mm, apiculate. 
Petals obovate, c. 8 by 3.5-4 mm, pale violet, the 
notch shallow, c. 1 mm deep. Anthers 1.3-1.5 mm 
long; filaments of the longer stamens c. 2 mm, 
those of the shorter c. 1 mm. Style 3-3.5 mm. 
Stigma globose, c. 1.5 mm thick, surrounded by 
the anthers at anthesis. Capsule densely strigulose 
with an admixture of erect, glandular trichomes, 
erect, 5-9.5 cm long, on a pedicel 1-2 cm. Seeds 
1.2 by 0.6 mm, coarsely papillose, dark brown, the 
coma 6-7 mm long, white, tinged with brown at the 
base. 

Distr. Continental SE. Asia (W. Nepal to 
Yunnan, south to the Khasya & Naga Hills), in 
Malesia: Central W. Sumatra (Mt Kerintji), one 
collection. 

Ecol. Along river-bank, c. 2000 m. 

4. Epilobium hirtigerum A. Cunn. Ann. Mag. Nat. 
Hist. 3 (1839) 33; Haussknecht, Monogr. Epilob. 
(1884) 291; H. Leveille, Ic. Gen. Epilob. (1910) 
t. 18; Allan, Fl. New Zeal. 1 (1961) 279; Bur- 
bidge & Gray, Fl. A.C.T. (1970) 276; Willis 
Handb. PI. Vict. 2 (1972) 464; Raven, D.S.I.R. 
New Zeal. Bull. 216 (1976) 141, f. 63-64, 65-66 
(maps). — E. brasiliense Haussknecht, Oest. 
Bot. Z. 29 (1879) 119. — E. sarmentaceum (non 
Haussknecht) Koord. Exk. Fl. Java 2 (1912) 704; 
Back. & Bakh. /. Fl. Java 1 (1963) 262. — 
E. cinereum (non A. Rich.) Raven, Blumea 15 
(1967) 273, pro specim. mal. 

Robust, erect, clumped perennial herb, 15-100 
cm, the underground parts not scaly ; plants densely 
strigulose, the stems pubescent all around. Lower- 
most leaves opposite, the rest alternate, densely 
strigulose, especially below and along the veins, 
nerves evident, narrowly lanceolate, 0.6-3 by 
0.2-0.5 cm, coarsely serrate, subsessile. Inflores- 
cence erect. Flowers erect, borne in the axils of 
upper leaves. Floral tube c. 1 mm across, c. 1 mm 
deep. Sepals 3-5.5 by 0.8-1.6 mm. Petals obovate, 
3-5 by 2-3.2 mm, white or very pale pink, the 
notch c. 1 mm deep. Anthers 0.7-1 mm long; 



1977] 



Onagraceae (Raven) 



111 



filaments of the longer stamens 2-2.8 mm, those of 
the shorter 1-1.8 mm. Style 1-3 mm. Stigma 
clavate, 1.5-2.7 mm high, 1-1.5 mm thick, sur- 
rounded by the anthers of anthesis. Capsule 
3-5(-6) cm long, on a pedicel 0-2 cm. Seeds 
0.9-1.2 mm long, 0.35-0.45 mm thick, finely 
papillose, brown, not beaked, obovoid, blunt at 
both ends, the coma 5-8 mm long, white. 

Gametic chromosome number, n = 18. 

Distr. Very widely distributed, South America 
(Argentina, Uruguay, Brazil: Santa Catarina), 
New Zealand, Australia, and Malesia: Lesser 
Sunda Is. (Timor, Sumbawa, Lombok) and East 
Java (Mt Tengger). 

Ecol. Moist places, grasslands; in E. Java the 
only locality is near the single small well on the 
otherwise dry and barren extinct volcanic cone of 
Mt Widodaren on Mt Tengger caldera, at 2100 m. 
In Lombok in Casuarina forest; in Timor in 
Eucalypt savannah; 1800-2200 m. Fl. Oct.-Jan. 

Note. Variable species. The Malesian specimens 
belong to a highly autogamous Australian form 
with small white flowers which occurs in swampy 
places throughout the lowlands of Australia (from 
Queensland to temperate W. Australia and Tas- 
mania) and is predominant in New Zealand. I have 
assumed that it came there from Australia and has 
spread again from New Zealand to South America, 
either by natural dispersal or by man. 

5. Epilobium kevsseri Diels, Bot. Jahrb. 62 (1929) 
486; Hoogl. Blumea Suppl. 4 (1958) 223; Borg- 
mann, Z. Bot. 52 (1964) 124, 143; Raven, Blumea 
15(1967)274, f. 1 (map). 

Clumped erect perennial herb 12-60 cm, often 
ericoid in aspect, ± woody at the base, the under- 
ground parts not scaly; plants finely glandular- 
pubescent. Lowermost leaves opposite, the rest 
alternate, coriaceous, subglabrous, the nerves 
obscure, margin revolute, narrowly elliptic to 
elliptic, 0.5-1.3 by 0.1-0.2(-0.4) cm, with a few 
coarse teeth on each side, subsessile. Inflorescence 
erect. Flowers erect, borne in axils of upper leaves. 
Floral tube 1-2 mm across, 0.7-1 mm deep. Sepals 
3-5.6 by 1-2.5 mm. Petals obovate, 4.5-8 by 2.8-4 
mm, purplish rose, the notch 1.5-2 mm deep. 
Anthers 0.7-1 mm long; filaments of the longer 
stamens 1.8-2.5 mm, those of the shorter 0.5-1.5 
mm. Style 2.5-4 mm. Stigma clavate, 1.5-2 mm 
high, 0.5-0.7 mm thick, surrounded by the anthers 
at anthesis. Capsule 3-6 cm long, glabrescent, on a 
pedicel 1-2.5 cm. Seeds 0.9-1.2 by 0.4-0.6 mm, 
papillose, brown, with a short pellucid beak, the 
coma 5-7 mm long, white. 

Gametic chromosome number, n = 18. 

Distr. Malesia: New Guinea (Mt Wilhelmina 
in West, common on many summits in East), many 
collections. Fig. 8. 

Ecol. Subalpine and alpine meadows and grass- 
lands, open shrubberies, swampy treefern grass- 
land, occasionally epiphytic on treefern trunks. 
Men in succession after ground-fires, 

(1600 )1950 3800 m.Ft. (Ian. (June Augjj Dec). 

Vern. Aingum, Tomba, gonema, C'himbu, 
yandepai, I nga lang., Wabag, papai, Enga lang., 
ttimitan, Mcndi lang., (iihmc. 
Notes. A distinctive species hut clearly belong- 
ing to the Australasian asscmbl 

plants, vvith their narrow leaves, are dis- 




Fig. 8. Range of Epilobium keysseri Diels (after 
Raven, 1967). 

tinctly ericoid in appearance, and thus parallel 
representatives of many other typically non-ericoid 
groups that occur in alpine and subalpine regions 
of New Guinea. 

6. Epilobium detznerianum Schltr ex Diels, Bot. 
Jahrb. 62 (1929) 485 ; Raven, Blumea 15 (1967) 277, 
f. 3, 5 (map). — E. papuanum Ridl. var. alpestre 
Ridl. Trans. Linn. Soc. Bot. II, 9 (1916) 58. — 
E. papuanum (non Ridl.) Hoogl. Blumea Suppl. 4 
(1958)228. — Fig. 7d. 

Clumped perennial herb 3-15 cm, the under- 
ground stems not scaly; plants mostly glabrous, 
with elevated, strigulose lines running down from 
the margins of the petioles and glabrous ridges 
running down from back of petioles. Leaves mostly 
opposite, alternate in the inflorescence, coriaceous, 
nerves obscure, broadly elliptic or ovate, obtuse at 
apex and base, entire, 0.3-1 by 0.2-0.7 cm; 
petiole 1-1.5 mm, short but distinct. Flowers 
nodding, the ovaries erect, borne in the axils of 
upper leaves. Floral tube 1.5-3 mm across and 
about as deep. Sepals 4.5-7 by 1-2.5 mm. Petals 
obovate, 7-14 by 3-6 mm, bright purplish rose, 
the notch c. 2 mm deep. Anthers 0.8-1.2 mm long; 
filaments of the longer stamens 4-5.5 mm, those of 
the shorter 3.3-4.5 mm. Style 5.5-8 mm. Stigma 
broadly clavate, 1.3-1.7 mm high, c. 1 mm thick, 
surrounded by or held just above the anthers at 
anthesis. Capsule erect, subglabrous, 4-5 cm long, 
on a pedicel 2.5-8.5 cm. Seeds (0.9— )I — 1 .5 by 
0.5-0.7 mm, not beaked, finely papillose, pale 
brown, the coma c. 8 mm long. 

Gametic chromosome number, n = 18. 

Distr. Malesia: New Guinea (Mts Carstensz & 
Wilhelmina in West, Telefomin, Mts Sarawaket & 
Wilhelm in East). Fig. 9. 





• 


% 


• 


4. 








X \^ i 








a. 





Fig. 9. Range of EpUohmm detxnerkuuan Schltr 
f.x Dims (after Raven, 1967). 



112 



Flora Malesiana 



[ser. I, vol. 8 2 



Ecol. Subalpine and alpine grasslands and 
ridges, open slopes and bogs, earth screes, sub- 
alpine forest and its grassy edges, rock clefts, cliff 
crevices in alpine thickets, on dripping slate land- 
slides, more rarely on sandy or gravelly gully 
beds; 3000-4500 m. Fl. (Jan.-Febr.-)May-Sept. 
(-Dec). 

Note. An attractive alpine species reaching the 
highest elevations in the genus in New Guinea. It is 
usually more condensed in habit than E. hooglandii 
and differs at once from that species in the glabrous 
ridge decurrent from the center of each petiole and 
the elevated pubescent lines decurrent from the 
edges of the petiole. It grows sympatrically with 
E. hooglandii and E. keysseri, but no intermediates 
have been observed; intermediates with the former 
species would be very difficult to detect. 

7. Epilobium hooglandii Raven, Blumea 15 (1976) 
278, f. 2, 6 (map). — E. pedunculare (non A. Cunn.) 
F.v.M. Trans. R. Soc. Vict. I, 2 (1889) 7. — E. 
detznerianum {non Schltr ex Diels) Hoogl. 
Blumea Suppl. 4 (1958) 228. — Fig. 7a-c. 

Caespitose perennial herb with decumbent 
branches, the erect portions 10-25(-45) cm long; 
plants glandular pubescent along elevated lines 
running down from the margins of the petioles 
below, more densely and uniformly so above, and 
with an admixture of strigulose pubescence in the 
inflorescence. Leaves mostly opposite, alternate in 
the inflorescence, subcoriaceous, nerves ± visible 
in dried material, broadly elliptic to ovate, acute or 
obtuse at apex and base, entire or with a few teeth 
on the margins, 0.5-1.3 by 0.3-0.9 cm; petiole 
1-3 mm, short but distinct. Flowers nodding, the 
ovaries erect, borne in the axils of upper leaves. 
Floral tube 1.4-2 mm across, 1-1.2 mm deep. 
Sepals 3-4 by 1-1.6 mm. Petals 6-8.5 by 2.5-4.5 
mm, pink to purplish rose, the notch c. 1.5 mm 
deep. Anthers 0.7-1 mm long; filaments of the 
longer stamens 3-6 mm, those of the shorter 2-4 
mm. Style 2.5-6 mm. Stigma clavate, 2-2.5 mm 
long, 1-1.2 mm thick, surrounded by the anthers 
at anthesis. Capsule erect, glabrescent, 5-8 cm long, 
on a pedicel 4-12 cm. Seeds 1-1.4 by 0.3-0.45 mm, 
finely papillose, brown ; coma 5-8 mm long, white. 

Gametic chromosome number, n — 18. 

Distr. Malesia: New Guinea (Mt Wilhelmina 
in West, many localities in East). 

Ecol. Subalpine and alpine meadows, fire- 
induced alpine grassland, near boulders, in alpine 
shrubbery, on peaty grassland, near waterfalls, in 
secondary forest on limestone cliffs, on stream- 
bank gravel, stony creek beds, along forest tracks, 
between grass tussocks in old lake basin, in pen- 
dent masses on rocks, occasionally in moist, 
forested areas; 2000-4150 m. Fl. May-Nov. 
(-Jan.). 

Vern. Dirimpia, Chimbu, Masul, nonami, 
Mairi, Mondo. 

Note. Differs from the closely related E. 
detznerianum by its evenly pubescent stems, and 



from E. prostratum by large flowers and seeds. It 
grows sympatrically with the other three New 
Guinean species. Occasional collections are inter- 
mediate with E. prostratum and obviously result 
from incidental hybridization. However, the two 
species keep in general amply distinct and are as 
well differentiated as most recognized species of 
the genus. 

8. Epilobium prostratum Warb. Bot. Jahrb. 16 
(1893) 15, 23; Raven, Blumea 15 (1967) 280, f. 4, 
7 (map). — E. papuanum Ridl. Trans. Linn. Soc. 
Bot. II, 9(1916)57. — Fig. 7e. 

Similar to E. hooglandii, but differing as follows: 
habit lax, spreading. Leaves 0.4-0.8 by 0.2-0.4 cm. 
Floral tube 0.8-1.1 mm across, 0.6-0.9 mm deep. 
Sepals 1.5-3 by 0.6-1.4 mm. Petals 2.5-5(-6) by 
1.3-2.5 mm, very pale pink to purplish rose, the 
notch c. 1 mm deep. Anthers 0.5-0.7 mm long; 
filaments 1.8-3.5 mm. Stigma clavate, 0.8-1.5 mm 
long, 0.5-0.9 mm thick, surrounded by the anthers 
at anthesis. Capsule 3.5-5 cm long, on a pedicel 
5-10 cm. Seeds 0.7-0.9 by c. 0.3-0.4 mm, finely 
papillose, brown, the coma 5-8 mm long. 

Gametic chromosome number, n = 18. 

Distr. Malesia: SW. Central Celebes (Latimod- 
jong Mts), Moluccas (Ceram), and throughout 
New Guinea. Fig. 10. 




Fig. 10. Range of Epilobium prostratum Warb. 



Ecol. Moist open places, especially on rocky 
alluvium bordering streams, stream-banks across 
treefern grassland, along river-bank in limestone 
scree, on fallen trees on open ridge, rarely in 
montane forest, sometimes colonizing landslips; 
( 1200-)1 900-3400 m, in Celebes and Ceram at 
c. 2750-3400 m. Fl. Jan.-Dec. 

Warburg's type specimen was found by Hell- 
wig at an exceptional low altitude of 1200 m in a 
streambed. 

Vern. New Guinea: kimbil, Enga lang., Poio, 
dirimpia, Chimbu, Masul, kokorabadi, Mandi lang. 

Notes. Closely similar to E. hooglandii, but 
readily distinguished for the most part by smaller 
flowers and seeds. Occasionally intermediate 
specimens (obviously hybrids) are found where the 
two occur together. 

In New Guinea it descends to lower altitudes and 
has correspondingly a much wider range. 



Introduced 

Fuchsia boliviana Carr. Rev. Hort. (1876) 150; as F. corymbiflora (non R. & P.) Back. & Bakh./. Fl. 
Java 1 (1963) 264. 

Locally cultivated and perhaps established in the mountains of West and East Java (Mts Malabar and 
Tengger) between 1500 and 2000 m. 



1977] Onagraceae (Raven) 113 

Easily distinguished from the following species by its drooping inflorescences of bright red flowers with 
a floral tube 5-6 (instead of less than 1) cm long. 

Fuchsia magellanica Lamk var. gracilis (Nich.) Bailey: Back. & Bakh. /. Fl. Java 1 (1963) 264; as 
F. coccinea (non Sol. ex Aiton) Curt. — Bunnemeijer, Trop. Natuur 10 (1921) 56, fig.; Wisse, ibid. 11 
(1922) 480, fig.; Hochr. Candollea 2 (1925) 480; Back. & Bakh./. Fl. Java 1 (1963) 264. 

This species, native of temperate South America, has repeatedly been reported to occur cultivated but 
also naturalized in anthropogenous places and on volcanic ash on the mountains of West, Central, and 
East Java (Mts Patuha, Malabar, Dieng, Sindoro, and Tengger) between 1000 and 2100 m, and in the 
mountains of Sumatra (Karo Lands). 

Oenothera stricta Ledeb. ex Link, En. Hort. Berol. 1 (1822) 377, ssp. stricta; as O. erythrosepala {non 
Borbas) Back. & Bakh./. Fl. Java 1 (1963) 262. — Oenothera sp. Doct.v.Leeuwen, Verh. Kon. Ak. Wet. 
A'damH, 31 (1933) 191. 

Docters van Leeuwen I.e. introduced this species from seed he collected in Hawaii and sowed on the 
summit of Mt Pangrango in West Java, at 3000 m altitude, in 1921. It is maintaining itself there and, 
although self-pollinating, is visited by Bombns rufipes (Heide, Dansk Bot. Ark. 5, 1927, 18) and doubtless 
by nocturnal insects as well. It is a native of temperate South America, widely cultivated and naturalized. 

It may be distinguished from all other species of the family in Malesia by its combination of a long 
floral tube and yellow petals; the flowers open at sunset. 

Cultivated 

Representatives of several genera — Clarkia and Gaura among them — are cultivated, mainly in the 
mountains. Backer & Bakhuizen van den Brink/, Fl. Java 1 (1963) gave an account of these. 



BIGNONIACEAE (C. G. G. J. van Steenis, Leyden) 

Trees, shrubs, lianas, very rarely herbaceous (extra-Mai); twigs often lenticellate 
and nodes with gland fields; spines very rare (extra-Mai). Stipules absent. Leaves 
simple or mostly compound (digitate or impari-l-4-pinnate), (in Mai.) decussate, 
rarely in whorls of 3^, often provided with glands underneath, in the New World 
often provided with terminal tendrils, rarely scattered or in pseudo-whorls (extra- 
Mai); domatia sometimes present (fig. 8b, 23h). Inflorescences bracteate, cymose, 
but not rarely thyrses contracted to racemiform or racemose inflorescences, or 
even reduced to solitary flowers (extra-Mai), terminal, axillary or from the old 
wood. Pedicels mostly with 1-2 bracteoles. Flowers usually very showy, rather large, 
bisexual, articulate with the pedicel or not. Calyx connate, closed in bud and later 
(not rarely irregularly) splitting into lobes, or cupular, or spathaceous, or lobed 
from the beginning and with equal or unequal, valvate lobes, developing earlier 
than the corolla, often glandular outside and inside with water and slime producing 
glands and hydathodes, persistent or circumscissile caducous along an abscission 
line. Corolla sympetalous, campanulate, tubular, funnel- or salver-shaped, mostly 
zygomorphic, lobes equal or unequal, valvate or imbricate in bud, tube often with a 
narrow cylindrical (constricted) lower part (basal tube) and a widened upper part 
(upper tube). Stamens 5 almost equal, or mostly 4 didynamous, the 5th sterile, 
rudimentary, adnate to the corolla tube, mostly inserted at the rim of the basal 
tube and not rarely (glandular) hairy at the insertion, more rarely inserted higher 
up. Anthers basifixed, 2-celled, rarely one cell barren or 1 -celled, introrse, dehiscing 
lengthwise, usually the anthers connivent in pairs ; anther cells often free and diver- 
gent, connective not rarely produced. Disk intrastaminal, mostly annular, rarely 
absent. Ovary superior, 2-celled, rarely 1- or 4-celled (extra-Mai) ; style filiform, 
stigma usually 2-lipped, sensitive. Ovules (in Mai.) in each cell on the septum in 
two or more rows of 3-cvd, mostly on 2 placentas. Capsule 2-valved, either loculicid 
with the septum perpendicular to the valves — sometimes provided with an addi- 
tional transverse false septum — or septicid with the septum parallel with the valves, 
or (extra-Mai) an indehiscent, 1 -celled, soft or hard-shelled, pulpy berry. Seeds in 
each cell attached to the dissepiment in one or more rows, inserted transverse to 
axis of fruit, anatropous, mostly on both sides with hyaline wings ; embryo exal- 
buminous, the cotyledons mostly notched, sometimes on both sides. Germination 
always epigeal. 

Distribution. About 120 genera and some 650 spp., mainly in the tropics and subtropics, 
roughly between 40° N and 30-35° S, very few in the warm-temperate zone; in Malesia: 14 
native genera of which 2 are endemic, viz Hieris in Penang and Lamiodendron in Papuasia. 
Among the remaining 12 one occurs through the Old World (Dolichandrone), 7 are shared with 
continental SE. Asia (two of which extend also to Africa and Madagascar: Fernandoa, Stereo- 
spermum) and 4 with Australia and Melanesia; the latter occur in Malesia only in the east except 
Deplanchea which ranges westward to Sumatra. 

In the family tropical Asia and Africa share a few genera {Markhamia, Fernandoa, Stereosper- 
mum, and Dolichandrone), but Africa and America share only one, viz Tecoma. This latter 
affinity goes further, though very disjunct via Campsidium (Chile) and Campsis (N. America and 
E. Asia) to Tecomanthe-Pandorea-Neosepicaea (Moluccas to Three Kings Is. and E. Australia). 
Otherwise there appear to be only two other transoceanic ties, viz tribe Crescentieae which is 
shared by Africa and the Americas, and the genus Catalpa which occurs in E. Asia and the 
Caribbean area. 

As Gentry (Brittonia 25, 1973, 227-230) has shown, the average number of species per genus 

(114) 



1977] Bignoniaceae (van Steenis) 115 

is only 5, which is very small in comparison with many other families, but can only partly be 
explained by a possibly small generic concept. There are quite a number of monotypic genera 
(in Malesia 5), but they are well defined in many characters and stand very apart. 

This, and the worldwide distribution of the family, and the disjunctions in ranges, definitely 
point to relict survival and ancient origin, onwards of which period the three tribes have under- 
gone a separate, independent development on the continents, mainly leading to differentiation in 
Indo-Australia and in the New World, with the greatest abundance in the latter. Unfortunately, 
the fossil evidence (only Tertiary) is meagre and untrustworthy (Schumann in E. & P. Nat. Pfl. 
Fam. 4, 3b, 1894, 208), both to macrofossils and to pollen. 

Ecology. Within the family there is a fairly wide coverage of habitats and there are quite a few 
which are confined to arid conditions (Rhigozum and Catophractes in S. Africa, Tecomella in 
Arabia, Phyllarthron bernierianum in Madagascar, Dolichandrone filiformis, D. heterophylla and 
the linear-leaved drought form of Pandorea pandorana in Australia). A few are rheophytic 
(Astianthus and Chilopsis in the Americas). A few are warm-temperate (Catalpa) or subtropical- 
alpine (Incarvillea incl. Amphicome in the Sino-Himalayas and Argylia and Campsidium in the 
Andes). One species is bound to the mangrove {Dolichandrone spathacea in Indo-Melanesia). 

The majority, however, belongs to the tropical forest, mostly the everwet type, but a fair num- 
ber in the seasonal type, below c. 2000 m; only Tecomanthe ascending to c. 3100 m in New 
Guinea. 

In Malesia most are evergreen, but Oroxylum indicum (fig. 5) and Dolichandrone spathacea can 
stand leafless in the dry season for many months. Fernandoa macroloba is also deciduous, as are 
the species of Stereospermum. 

Habit. The majority of the Malesian genera are small or large trees, the only climbing genera 
being Nyctocalos, Hieris, Tecomanthe, Pandorea, and Neosepicaea; in subalpine heathland 
Tecomanthe may be forced to creep on other vegetation. 

Most trees are of medium size, but species of Stereospermum, Fernandoa, Pajanelia, and also 
Radermachera gigantea may attain quite good dimensions. Oroxylum is a short-lived nomad tree. 

Dominance. Almost all species occur scattered in the forest and several are very rare indeed. 
An exception is Dolichandrone spathacea, bound to the swampy, brackish inner mangrove, 
which according to Corner (Wayside Trees, 1940, 164) is in the North of the Malay Peninsula, 
in Perlis, a feature of the country, flanking roads and standing as an upright poplar in the rice- 
fields. Fig. 16. In Great Natuna I. (NW. off Sarawak) I found Pajanelia longifolia locally very 
common in coastal forest, but this was probably encouraged by devastation. In secondary forests, 
on earth slides, abandoned fields, and on fresh volcanic ash Radermachera glandulosa and R. 
gigantea may be frequent in the pioneer upgrowth in Java (also on Krakatao), but this high 
frequency is local ; Koorders mentioned it for G. Telemojo and Pringombo in Central Java. Even 
Oroxylum indicum which is a nomad plant bound to secondary growths is always found in only 
a few specimens. Lamiodendron magnificum was once mentioned by Brass as forming a com- 
munity on a gravel bank behind the beach, in Normanby I., but this tree is extremely rare other- 
wise. 

Flower biology. In many species (like in several Gesneriaceae, Verbenaceae, Solanaceae) the 
calyx develops much earlier than the corolla and is closed in bud. Inside of the calyx with water 
and slime producing glands and hydathodes in which the corolla develops. These so-called 
waterbuds are very characteristic, especially in such large-flowered species as Spathodea campanu- 
lata, the tulip tree, which derives a Dutch and Malay name from this feature (spuitjesboom, 
panchot) which is enjoyed by children to play with. It is one of the few biological phenomena 
which are entirely confined to the tropics, as far as I am aware. 

Treub (Ann. Jard. Bot. Btzg 8, 1889, 38-46, t. 13 15) made an anatomical study of the glands 
inside the calyx of Spathodea campanulata and on his instigation ( rR] simi i examined the exudate, 
dissolved organic and inorganic substances, which appeared similar to those of Icaf-hydaihodcs. 
Koordiks tihid. 14, 1897, 354 469, t. 21-27) extended this subject with research on some other 
genera of Bignoniaeetu (Parmenttera, Klgelta, Crescentia, Fernandoa adenophylla, Radermachera 
gigantea) and some other plants, confirming Tkm B'l icsults. 

Pollination I lower-shape, -colour, -position, anil -scent arc very different in the mostly showy 
flowers of Bignomai car, and the syndromes attract different visitors 



116 Flora Malesiana [ser. I, vol. 8 2 



Bats are frequently visiting species of certain genera, another phenomenon restricted to tropi- 
cal plants. According to Faegri & van der Pijl (Principles Pollination Ecology, ed. 2, 1971, 
154) the attraction syndrome is: nocturnal anthesis, whitish, creamish or drab greenish or dark 
purple colour, stale or sour, unpleasant smell reminiscent of fermentation at night, large quantity 
of nectar and pollen in large anthers, large-mouthed and coarse flowers on strong stalks sticking 
out of the foliage or caulifiorous to fiagelliflorous flowers, thus coming into easy reach for landing. 
This is found in Malesia in several cultivated genera (Kigelia, Crescentia, Parmentiera, Mark- 
hamia, etc.) but occurs also in the native Fernandoa adetwphylla, Pajanelia, and Oroxylum. 
Fig. 7. Notwithstanding the many papers and records of observation — corollas show claw marks 
after these visits — it is not proved to my satisfaction that visits of bats are compulsory for polli- 
nation cq. fertilisation, experimenting being in this field deplorably meagre. My doubt is streng- 
thened by observations by Harris & Baker in Ghana where Kigelia is native (J. West Afr. Sc. 
Assoc. 4, 1958, 28) and can set fruit in absence of bats; they observed also frequent visits by 
sphingids but they doubt effective pollination by these. 

Birds, humming birds and sun-birds, frequently visit certain species, the attraction syndrome 
being: tubed, vividly coloured (orange, scarlet), diurnal, mostly odorless, nectar-producing 
tubular flowers. Here also many observations are made, e.g. in Tecoma (Tecomaria). To this class 
belong in Malesia some species of Radermaehera (R. rami flora), Neosepicaea, Tecomanthe, and it 
can be expected for Deplanchea. Also the cultivated Spathodea campanulata is frequented by 
birds (cf. Beumee, Trop. Natuur 14, 1925, 28-30), notably kutilans and ?djalaks, at Bogor; they 
severely damage the corolla. Here again the question whether bird-visits are compulsory for 
pollination cq. fertilisation is inadequately supported by experiments. Caution is necessary to 
conclude to the necessity of cross-pollination, as e.g. Hunter (Rec. Auckl. Inst. Mus. 6, 1967, 
169-170, t. 24) recorded that in Tecomanthe speciosa, of which cuttings of a single plant led to its 
cultivation, fertilisation — that is selfing — could be effected by hand-pollination, but later also 
naturally by bees, although far from its native habitat. 

Moths. A few species have the moth-attraction syndrome for flower visitors, which implies: 
nocturnal, very fragrant flowers with abundant honey, in mostly pale or white, long-tubed or 
salver-shaped corollas. Fig. 1, 8, 15. This is found in species of Nyctocalos, Dolichandrone, Hieris, 
Millingtonia, and some species of Radermaehera (e.g. the Chinese R. sinica, R. pentandra, and 
R. peninsular is). Probably long-tongued sphingids (hawk-moths) visit these flowers. 

Bees and butterflies. Possibly bees visit flowers of species not belonging to the three categories 
mentioned above. 

The ecological role of the many sorts of extra-floral nectaries in unexplained. 

It is a fact that in general fruit setting is scarce in Bignoniaceae; in several the fruit was only 
occasionally found long after the plant had been described in flower. In Malesia the fruit is still 
unknown of Hieris and Lamiodendron. Even after abundant flowering fruit production is often 
very low with 1 or 2 fruits in each inflorescence, except in Radermaehera glandulosa and Tecoma 
stans. With all these flower visit devices one would expect otherwise. 

Dispersal. Bignoniaceae occur throughout the tropics and several are still found in the sub- 
tropics of the whole world. One might ascribe this to their having winged seed (except Crescentieae 
and a few other exceptions), but against expectations they are almost absent from oceanic 
islands. Bignoniaceae occur all along the coasts of the West Pacific, notably in New Guinea and 
in Australia species of Tecomanthe and Pandorea are not rare, but the only occurrence in the West 
Pacific islands is a common Australian Pandorea in New Caledonia and Lord Howe I., and a 
peculiar Tecomanthe in a single locality of the Three Kings Is., the northernmost territory of New 
Zealand. 

Obviously wind dispersal has not been as effective as one would expect. 

Dispersal by seawater is common in Dolichandrone spathacea, a back-mangrove species, 
ranging from the western Deccan Peninsula to North Luzon, south to Timor and southeastwards 
to New Caledonia; the range is almost continuous without gaps. Fig. 14. It is most peculiar, 
however, that so far it has never been found in the mangroves of northern Australia. Its seeds have 
thickish corky wings instead of flimsy wings as usual in most members of the family (except the 
fleshy indehiscent fruits of the Crescentieae and a few other exceptions as e.g. Pauldopia) and are 
most excellently adapted to be dispersed by seawater. 



1977] Bignoniaceae (van Steenis) 117 

Seedlings. These are very uniform in all tribes of the family; in the embryo the foliaceous 
cotyledons are flat in one plane, mostly emarginate at both ends, hypocotyle and rootlet are 
small. Germination is epigeal by stretching of the hypocotyle. The first leaves are mostly simple, 
as in most compound-leaved families; they are often dentate. In a very few genera with thick seeds 
there may develop — possibly by intrusion of the testa — a false septum in the seed, in Malesia 
notably in Stereospermum and here also the cotyledons are folded. The only exception is men- 
tioned by Lubbock (Contr. Knowl. Seedlings 2, 1892, 334, fig. 569) for 'Bignonia insignis' with 
fleshy connate cotyledons and hypogeal germination; the name is evasive and at Kew the identity 
could not be traced ; presumably the record rests on an error. 

Literature: E. Bureau, Monogr. Bign. (1864); J. Lubbock, Contr. Knowl. Seedlings 2 (1892) 
332-345, fig. 569-575; R. S. Troup, Silvic. Indian Trees 2 (1921) 684-693; J. A. Duke, Ann. Mo. 
Bot. Gard. 52 (1965) 349, pi. 20; G. de la Mensbruge, La germination et les plantules essences 
arbres Cote d'lvoire; Techn. For. Trop. Nogent-sur-Marne, Paris (1966) 332-333; D. Burger, 
Seedlings (1972) 52-54; C. S. Schopmeijer, Seeds Woody PI. U.S., Agric. Handb. 450 (1974) 260, 
281, 321. 

Juvenile plants of Pandorea pandorana show leaves very different from the mature foliage, in 
having many jugae and being coarsely dentate. Tecoma filicifolia Nichols, was based on such 
material. This led also to a serious misinterpretation of Tecoma leptophylla Bl., from New 
Guinea, of which the juvenile leaves (fig. 37b) are Pandorea pandorana but the flowers belong to 
Neosepicaea. 

Taxonomy. Since the basic work on the systematy by E. Bureau (Monographic, 1864), the 
treatment of the family in Flora Brasiliensis by Bureau & Schumann (1896-97), and the treat- 
ment of Schumann in the Pflanzenfamilien (1895) the traditional subdivision of the family in 5 
tribes has proved satisfactory. Crescentieae with 1 -celled berries occur in Africa and the Americas, 
two other monogeneric tribes are South American, while the bulk of the family belongs to Big- 
nonieae and Tecomeae, of which the latter are about balanced as to number of genera in the Old 
and New World, but Bignonieae are predominantly American. These two tribes are largely dis- 
tinguished on the dehiscence of the fruit, loculicid in Bignonieae and septicid in Tecomeae. 

In passing it may be remarked that Gentry (PI. Syst. Evol. p. 126, 255) recently advocated that 
Crescentieae of the neo-tropics and of Africa-Madagascar are of separate descent and would 
represent two parallel evolutionary lineages; this suggestion is more based on geographic argu- 
ment and evolutionary hypotheses than on morphological arguments. 

The delimitation against other families of Sympetalae is well-defined, but there are a few genera, 
notably Wightia and Paulownia, which are sometimes referred to Bignoniaceae, though Fenzl 
(Denkschr. K. Bay. Bot. Ges. Regensburg 3, 1841, 227-230), Bureau, Schumann, von Wett- 
stein, and other specialists referred them to Scrophulariaceae. A survey of opinions I gave in my 
paper on Wightia (Bull. Bot. Gard. Btzg III, 18, 1949, 214-216), in which I excluded it from 
Bignoniaceae. Even recently Paulownia is sometimes casually treated as Bignoniaceous {e.g. 
Schopmeijer, Agric. Handb. U.S.A. 450, 1974, 527), although the embryo is embedded in 
endosperm; furthermore the stigma is different from that in Bignoniaceae, the anthers have no 
prolonged connective, there is no rudimentary stamen and the seeds are provided with several 
wings and seem to be laterally attached, not transverse as in Bignoniaceae. For Wightia I tabulated 
(I.e.) the relation to both families. Its seeds have no endosperm, but the absence of a staminode, 
the structure of the stigma, the central placenta and the absence of a produced connective on the 
anthers point distinctly to Scrophulariaceae. The seed is quite differently attached as compared 
with Bignoniaceae, viz laterally and the wing surrounds the entire seed. Its wood has two kinds 
of medullary rays, narrow and broad ones, a character which, at least in Malesian Bignoniaceae, 
is absent. 

Though the capsule in Wightia is septicid and in Paulownia loculicid, both genera have the 
same kind of axile placentation, in which the thickened placenta becomes detached from the 
valves as a subquadrangular seed-cake, showing their close affinity, completely differing from the 
situation in liignoniaceae. 

According to Suryakanta (J. Palyn.9, 1973, 73) the pollen of both genera differs from that in 
[iii'iuimiK riir and resembles that <>f SCTOphtllariacecU 

Nakai (J. Jap. Bot. 24, 1949, 13) accommodated Paulownia in Paulowniaceae, probably in- 



118 Flora Malesi ana [ser. I, vol. 8 2 

duced mostly by its arboreous habit and fruit; they certainly merit to be placed in a separate 
tribe or subtribe of Scrophulariaceae. We regard nowadays the arboreous habit as primitive in 
herbaceous families and we might conclude that they are ancient relicts from a period when 
Bignoniaceae and Scrophulariaceae had a common matrix. 

Also in South America there are two woody genera of the Scrophulariaceae which were at 
times referred to Bignoniaceae, viz Schlegelia (syn. Dermatocalyx) and Gibsoniothamnus, accord- 
ing to Gentry (Fieldiana, Bot. 34, 1971, 55; Ann. Mo. Bot. Gard. 61, 1974, 533-537); see also 
Leinfellner (Oest. Bot. Z. 121, 1973, 13-22). They are (hemi-?) epiphytic shrubs or lianas, a 
similar habit as in Wightia. 

Genetics. Chromosomes. Darlington & Wylie (Chrom. Atlas, 1955) and Moore (ed.) 
(Regn. Veg. 90, 1973) gave for 26 genera x = 20 (2n = 40) and they belong to Tecomeae, 
Bignonieae and Crescentieae, both from the palaeo- or neotropics. There is one higher number 
x = 22 (Amphilophium, South America, Niedzwedzkia = Incarvillea) and several lower ones: 
Pandorea, and some doubtful countings in Tecoma x = 19, Tecomanthe dendrophila 2n = 36 
(Christine Brighton in litt.), Jacaranda x = 18, Tecoma capensis x = 17, Oroxylum, Milling- 
tonia, Argylia (from South America) x — 15, Spathodea x = 13, and Incarvillea x = 11. In 
supplement indices Campsis is also given as 16 and Oroxylum as 14. 

I have scanned the numbers of Scrophulariaceae, Gesneriaceae and Verbenaceae, but can find 
no reliable ties, Bignoniaceae being obviously more homogeneous than those. 

The number given for Paulownia, 2n = 40, x = 10, might as well fit Bignoniaceae as Scrophu- 
lariaceae. 

Hybridisation. Not many species hybrids are known to me, but those known are interesting, 
as there are at least two between species of East Asia and SE. North America which are now very 
disjunct after the Pleistocene Ice Age; it is not impossible that they formed part of more con- 
tinuous populations in the warmer Pliocene via Beringia. This idea is supported by the fact that 
in both cases the hybrids are fertile. 

E. C. Smith (J. Arn. Arb. 22, 1941, 219) reported on Catalpa ovata Don x C. bignonioides 
Walt. (= x C. syringifolia Sims). Haploid all have 20 chromosomes (Sax, J. Arn. Arb. 14, 1933, 
274). 

Then there is x Campsis tagliabuana (Viviani) Rehder, a hybrid between the Chinese C. 
grandiflora (Thunb.) K. Sch. (C. chinensis (Lamk) Voss.) and C. radicans (L.) Seem, which pro- 
duces fertile progeny (cf. Stearn, Bot. Mag. 169, 1953, t. 198). 

The third one is also bi-continental, Tecoma smithii W. Watson (Gard. Chron. 14, 1893, 649, 
fig. 104; E. Smith, ibid. 16, 1894, 64; cf. also Gartenflora 44, 1895, 51, fig. 14). This is a reputed 
hybrid, which E. Smith made at Adelaide, in 1882, between T. velutina (a hairy variety of T. stans) 
and T capensis. It was propagated by cuttings, but it produced seed and its offspring of seedlings 
diverged in size and flower colour. Curiously Sprague, in a succinct note (Fl. Cap. 4, 2, 1904, 
448) reduced it to T. alata DC, without referring to its hybrid nature. 

Anatomy. Wood. Of the Malesian Bignoniaceae only a small proportion of tree species is 
known wood-anatomically; the climbing and scandent species are fully unexplored in this 
respect. Anomalous structure has, however, been described for several genera outside Malesia. 
As far as known, the Malesian tree genera are wood-anatomically rather homogeneous: with 
simple, rarely also reticulate, perforations to the vessels, homogeneous rays, mainly paratracheal, 
aliform or confluent parenchyma, and fibres with simple to minutely bordered pits. Except for its 
unusually narrow rays, Dolichandrone spathacea from the mangrove swamps does not differ 
appreciably from the inland genera in its wood structure. 

Leaves. Very poorly known for the Malesian representatives. Diversity of stomatal type and 
indumentum (non-glandular and glandular hairs in a variety of forms) certainly deserves detailed 
comprehensive studies, which will probably yield important additional taxonomic characters. 

References: General surveys: Solereder, Syst. Anat. Dicot. Stuttgart (1899, 1908); Metcalfe 
& Chalk, Anat. Dicot. Oxford (1950). — Wood: Janssonius, Mikrographie des Holzes 4 
(1926) 721-753 (Dolichandrone, Oroxylum, Stereospermum); Panshin, Philip. J. Sc. 48 (1932) 
143-205 (Dolichandrone); Desch, Mai. For. Rec. 15 (1941) 50 (Deplanchea, Dolichandrone, 
Pajanelia, Stereospermum); Janssonius, Blumea 6 (1950) 450-452 (affinities); Sebastine, J. 
Ind. Bot. Soc. 34 (1955) 299-306 (Pajanelia); Jutte, Nova Guinea 10 (1959) 242 (Deplanchea). — 



1977] Bignoniaceae (van Steenis) 119 

Leaves: Siebert, Ann. Mo. Bot. Gard. 35 (1948) 123-136 (glands); Paliwal, Flora 159 (1970) 
124-132 (stomata). — P. Baas. 

Pollen morphology. Bignoniaceae have a long history of pollenmorphological study, 
starting with the pioneer studies of H. Mohl (1835). The first author to present a detailed pollen- 
morphological survey of the family, drawing attention to the taxonomical significance of the 
pollen characters, was Urban (1916). He concluded that (1) any attempt to base the main sub- 
division of the family on pollen characters would group together taxonomically unrelated genera, 
(2) for generic delimitation pollen had limited value. Later studies by several authors have 
confirmed this (cf. Buurman, in the press). 

Inaperturate, tricolpate, stephanocolpate and pericolpate apertural types occur in the family. 

In a few genera tetrads are found. 

Size ranges between 25 pun in Astianthus antisyphilitica and 100 pirn in Nyctocalos cuspidata, 
shape varies between suboblate and subprolate. 

Remarkable is the amount of variation which may occur within genera or even intraspecifically. 
In Stereospermum inaperturate, tricolpate and perisyncolpate pollen is found, while in Anemo- 
paegma longepetiolatum inaperturate, stephanocolpate, pericolpate grains as well as tetrads 
occur. In such cases sculpture affords more constant characters. 

The tricolpate type is dominant in the family and is found in all four tribes. It rarely shows well 
developed equatorial endoapertures. Instead, characteristically ruptured aperture membranes 
are present, especially in Tecomeae. Operculate colpi occur in Argylia. A subdivision of the tri- 
colpate type is possible on sculpture, which mostly varies between perforate and reticulate. 

In some genera very complex pollen grains are present, such as those of Nyctocalos (fig. 2). 

The tricolpate pollen grains in Bignoniaceae resemble those in Scrophulariaceae and Myopora- 
ceae. The similarities with Pedaliaceae, stressed by Erdtman (1952) refer to a rather specialized 
pollen type and may not reflect close affinity. 

References: Mohl, Ann. Sc. Nat. II, 3 (1835) 304-346; Urban, Ber. Deut. Bot. Ges. 3 I (1916) 
728-758; Erdtman, Pollen morphology and plant taxonomy. Angiosperms. Almqvist & 
Wiksell, Stockholm (1952) 73-74; Buurman, Contribution to the pollenmorphology of the 
Bignoniaceae, with special reference to the tricolpate type (in the press). — J. Muller. 

Chemotaxonomy. Bignoniaceae share a number of biochemical tendencies with 
Verbenaceae, Labiatae, Scrophulariaceae and with several other families of Wettstein's Tubi- 
ftorae. Most of their outstanding chemical characters were already mentioned and discussed in 
my 'Chemotaxonomie der Pflanzen' vol. 3 (1964) 268-281, 645-646, to which the reader is 
referred. Much phytochemical information, however, became available only in more recent time. 
Recent results confirm the trends already apparent in 1963; they are summarized in the following 
pages. Chemical characters of Bignoniaceae may ultimately prove to be very useful in tracing 
inter- and intrafamiliar relationships. 

( 1 ) Most members seem to produce and accumulate iridoid glucosides (formerly often called 
pseudoindicans). Since a long time Bignoniaceae are known to contain labile glycosidic bitter 
principles. Such a compound was isolated in 1888 from the bark and fruits of Catalpa bignonioides 
Walter and called catalpin (name changed later to catalposide). The structure of catalposide was 
definitely established in 1962; it is an aucubin-type (C g -aglucone) ester glucoside and one of the 
first pseudoindicans for which clearcut structural and biogenetic relationships with iridodial 
and nepentalactone were demonstrated (hence the name iridoid glucosides for a presently very 
large group of constituents of dicotyledonous plants). Catalposide (tastes bitter) is an ester of 
/7-hydroxybenzoic acid with catalpol (« 7,8-epoxy-aucubin). Catalpol and catalposide occur in 
all species of Catalpa (leaves, stems, fruits) and catalpol ( catalpinoside) was also isolated from 
barks of Paulownia tomentosa Steud. and P. fargesii Franch. where it occurs together with 
synngin (V. Plouvier, C. R. Ac. Sc. Paris 272D, 1971, 1443). Probably catalpol and catalposide 
occur in many more members of the family. In most recent times some related glucosides were 
isolated from Bignoniaceae. Vanilloyl-catalpol ( amphicoside) is a constituent of Amphicome 
emodi Lindl. and veratroyl-catalpol OOCUTI in TtCOHItUa umlulata Sum 5-Hydroxycatalpol 
| macfadyenosidc) was isolated from Macjadycna cynanchoides MORONO. All iridoid glucosides 
mentioned hitherto have structures based on the aucubin-dcmativc catalpol. I he first non- 
aucubin-typc glucoside described from Bignonim; ■</<■ is tccomosule with I C 10 -aglucone; it was 



120 Flora Malesiana [ser. I, vol. 8 2 

isolated from Tecoma capensis Lindl. (A. Bianco et al. Gazz. Chim. Ital. 105, 1975, 195). It is to 
be expected that much more iridoid glucosides will be detected in the family in future. 

(2) Some Bignoniaceae produce alkaloids. So far only pyridine-type and piperidine-type 
alkaloids with an iridoid C 10 - or rarely C 9 -skeleton were identified definitely in species belonging 
to this family. This fact strengthens the belief that the tendency to produce iridoid compounds is 
a very important character of Bignoniaceae. Thusfar simple iridoid alkaloids were described for 
species of Campsis (boschniakine), Incarvillea (plantagonine, indicain), Tecoma (tecomanine, 
tecostidine, tecostanine, boschniakine, 4-noractinidine and several derivatives of skytanthine). 
The basic constituents of Amphicome (now reduced to Incarvillea), Newbouldia and other genera 
may belong to the same group of alkaloids. 

A recent review of the chemistry, distribution and systematic meaning of all presently known 
main groups of iridoid plant constitutents was published by S. Rosendal Jensen et al. (Bot. 
Notis. 128, 1975, 148-180). 

(3) Many Bignoniaceae synthesize naphthaquinones and corresponding anthraquinones by 
prenylation of o-succinylbenzoic acid. This pathway to quinonoid naphthalene- and anthracene- 
type secondary metabolites is presently known from taxa belonging to Rubiaceae, Verbenaceae, 
Scrophulariaceae, Bignoniaceae and possibly Acanthaceae and Gesneriaceae. In roots, woods 
and barks of Bignoniaceae lapachol, lapachonone, a- and p-lapachone and dehydro-a-lapachone 
occur frequently. These monomeric naphthaquinonoid compounds are often accompanied and 
sometimes replaced by more complex dimeric constituents like tectol, guayacanine and guayine 
and by corresponding anthraquinones such as tectoquinone and 2-methyl-3-hydroxyanthra- 
quinone. Woods which contain appreciable amounts of these quinonoid compounds are more or 
less resistant to marine borers, white ants and Fungi. At the same time such woods may be the 
causes of skin irritations and of allergic skin diseases in man. Lapachol- and tectoquine-type 
substances are presently known from species of the genera Catalpa, Heterophragma, Kigelia, 
Paratecoma, Phyllarthron, Stereospermum, Tabebuia, Tecoma, Tecomella, and Zeyhera. R. H. 
Thomson has reviewed the chemistry and distribution of quinones and related compounds in his 
book 'Naturally occurring quinones' (2nd ed. 1971). The phthalide catalpalactone from the wood 
of Catalpa bignonioides Walter and C. ovata G. Don arises from the same pathway as lapachol 
and its congeners (H. Inouye et al. Chem. Pharm. Bull. Tokyo 23, 1975, 384, 392, 2523). On the 
other hand it should be stressed that the red-coloured naphthaquinones of Boraginaceae (e.g. 
alkannin) which are structurally very similar to lapachol are produced along a totally different 
biosynthetic pathway (cf. E. Leistner, Chinoide Farbstoffe, Ber. Deut. Bot. Ges. 88, 1975, 
163-178). 

(4) The "tannins" mentioned for many Bignoniaceae in the older phytochemical literature 
{e.g. Dekker, 1913) seem to be glycosides and esters of o-diphenolic compounds. Orobanchin 
(= verbascoside)-type ester glycosides were definitely demonstrated to occur in species of 
Campsis, Catalpa, Eccremocarpus and Pandorea. A review of this type of polyphenols plant 
constituents which simulate true tannins in some respects is to be found in my 'Chemotaxonomie 
der Pflanzen' vol. 5 (1969) 250-252. Orobanchin yields a molecule of caffeic acid, 3,4-dihydroxy- 
phenylethanol, glucose and rhamnose each. Just as in most other families of Sympetalae true 
tannins are replaced in Bignoniaceae by more or less complex esters and glycosides of o-diphenolic 
cinnamic acid derivatives. Moreover, simple esters of caffeic acid and biosynthetically related 
derivatives of cinnamic and benzoic acids are present in large amounts in many Bignoniaceae. 
The recent investigations of V. B. Pandey and B. Dasgupta with the bark of Tecomella undulata 
Seem, (veratroylglucose = tecomin: Experientia 26, 1970, 1187) and of M. Sugumaran et al. 
with leaves of 7Vcoma starts H.B.K. (16 aromatic acids: Ind. J. Exper. Biol. 13, 1975, 93) exemplify 
this trend. /7-Hydroxybenzoic acid is present as an ester in all species producing catalposide; 
probably this phenolic acid is rather ubiquitous in the family. The presence of appreciable 
amounts of hydroquinone (in living cells as the glucoside arbutin?) in leaves of Jacaranda 
mimosaefolia D. Don (S. Sankara Subramanian et al. Phytochemistry 12, 1973, 220) might be 
connected with a strong tendency to produce and accumulate />-hydroxybenzoic acid; if this is 
actually the case hydroquinone (and arbutin?) may be detected in much more Bignoniaceae in 
future. Jacaranone, a quinonoid compound which exhibits antitumor and cytotoxic activity was 
recently isolated from leaves and twigs of Jacaranda caucana Pittier (M. Ogura et al. Lloydia 



1977] Bignoniaceae (van Steenis) 121 

39, 1976, 255); it seems to be derived from tyrosine and is chemically very similar to the Cornus 
quinol glucoside (= cornoside) which is also present in leaves of Digitalis purpurea (Bot. Notis. 
128, 1975, 174). 

(5) According to J. B. Harborne (Phytochemistry 6, 1967, 1643) leaf flavonoid patterns of 
Bignoniaceae are close to those of Acanthaceae, Gesneriaceae, Labiatae and Scrophulariaceae. 
Features which support such a statement are the replacement of flavonols by flavones in many 
species, the relatively frequent occurrence of flavones with an unsubstituted B-ring {e.g. chryson, 
baicalein), of 6-hydroxylation of chrysin (baicalein), apigenin (scutellarein) and luteolin (6- 
hydroxyluteolin) and of O-methylation of flavones. The latter trend is illustrated by Zeyhera 
tuberculosa Bur. ex Verlot which contains 5,6,7-trimethoxyflavone and 5,6,7,8-tetramethoxy- 
flavone in leaves (J. P. Kutney & H. W. Hanssen, Phytochemistry 10, 1971, 3298). The bitter 
principle of the fruits of Sparattosperma vernicosum Bur. & K. Sch. was shown by J. P. Kutney 
et al. (Phytochemistry 9, 1970, 1877) to be the 7-neohesperidoside of pinocembrin (= 2,3- 
dihydrochryson). 

(6) Free triterpenic acids occur in appreciable amounts in leaf waxes of many families of 
Tubiflorae (especially Verbenaceae, Labiatae and Plantaginaceae) and related orders. It is of 
interest in this respect that ursolic acid was isolated in recent time from leaves of Bignonia 
diversifolia H.B.K., Campsis radicans Seem., Catalpa bignonioides Walter, Heterophragma 
quadriloculare K. Sch., Jacaranda mimosaefolia D. Don (not definitely identified) and Paulownia 
tomentosa Steud. The bark of Jacaranda mimosaefolia yielded lupenon. 

(7) Many members of Verbenaceae, Labiatae, Scrophulariaceae and Plantaginaceae replaced 
starch by stachyose-type oligosaccharides as storage carbohydrates. The same trend seems to 
exist in Bignoniaceae. Large amounts of stachyose occur in species of Catalpa (roots, wood, bark), 
Newbouldia laevis Seem, (roots) and Paulownia tomentosa (stem). 

(8) Most representatives of Tubiflorae produce starch-free seeds which are rich in proteins and 
oils. The seed oils are often characterized by a high degree of unsaturation. In this respect 
Bignoniaceae conform to the rule. Their seeds generally contain 20-35 % of oil. In some taxa 
oleic and (or) linolic and (or) linolenic acid are the only major fatty acids of the seed oils (e.g. 
species of Crescentia, Niedzwedzkia — Incarvillea, Paulownia and Stereospermum). In other taxa 
the 'normal' fatty acids are accompanied or replaced by large amounts of unusual fatty acids 
such as conjugated trienoic acids (species of Catalpa, Chilopsis, Jacaranda), C 26 -keto-acids 
(Cuspidaria pterocarpa DC), octadeca-rra«5-3,c/5-9,cw-12,cw-15-tetraenoic acid (Tecoma stans 
H.B.K..) or hexadec-9-enoic and octadec-11-enoic acid (Doxantha unguis-cati Miers). M.J. 
Chisholm and C. Y. Hopkins discussed the chemistry of seed oils of 1 1 species representing 4 
tribes (Canad. J. Chem. 43, 1965, 2566). 

The preceding phytochemical picture places Bignoniaceae phytochemically very close to a 
number of families of Tubiflorae, especially Verbenaceae, Labiatae and Scrophulariaceae. Still 
other constituents are known from Bignoniaceae. Lack of acquaintance with their structures and 
(or) with their distribution, however, does not yet allow a systematic evaluation. Saponins, which 
seem to be rather widespread in the family but were never investigated in detail, belong to these 
chemical characters. The same holds for a number of phenolic compounds isolated in recent 
time, such as the lignans sesamin and paulownin from Paulownia tomentosa Steud. and Phyl- 
larthron comorense DC, the dilignol (a lignan-type compound) zeyherol from Zeyhera digitalis 
Hoehne and the dihydroisocoumarins 6-methoxymellein, kigelin and 6-demethylkigelin from 
Kigelia pinnata DC. 

Concluding it may be stated that the intimate relationships between Bignoniaceae and Scrophu- 
lariaceae which are indicated by genera like Catalpa and Paulownia (often placed in Scrophu- 
lariaceae) are confirmed by phytochemistry. At the same time phytochemistry stresses a very 
close coherence of a core of families of Tubiflorae; this core comprises Scrophulariales sensu 
Cronquist (1968) and Lamiales sensu Takhtajan (1969). — R. Hignauer. 

Uses. There are no outstanding qualities marking Bignoniaceae as useful plants, otherwise 
than ornamentals and these concern mostly the introduced species for which I refer to the special 
key and account at the end. There are magnificent native species notably of Icconumthe but they 
have as yet not become in general use. 

Good roadside trees are Millingtonia hortensis and Spathodea campanulata. A highly esteemed 



122 Flora Malesiana [ser. I, vol. 8 2 

vegetable (lalab) with the Sundanese is Oroxylum indicum (flowers, buds, and very young pods). 

For re-afforestation and holding terraces on slopes the pioneer qualities of species of Rader- 
machera and Deplanchea might be useful. 

The timber is in general not valuable and in nature not available in sufficient quantity. The soft 
wood of Millingtonia hortensis was advertized as useful for tea-boxes. The only species yielding 
sizeable timber of good quality are: Fernandoa macroloba, Pajanelia longifolia, Radermachera 
gigantea, and the three species of Stereospermum, which all may be valuable for silviculture. 

Terminology. The shape of the corolla has been defined as tubular (fig. 4b, lOe) in which 
case there can be a distinction in a basal tube (on apex of which inside the stamens are inserted) 
as in fig. 4b; funnel-shaped (fig. 23d, 26c), hypocrateriform or salver-shaped (fig. la, 8c, 15) or 
infundibuliform (fig. 32a). 

Notes. Since my thesis (Rec. Trav. Bot. Neerl. 24, 1927, 787-1049), here always cited as 'Thesis 
(1927)', and subsequent revision in Bull. Jard. Bot. Btzg III, 10 (1928), I have remained always 
much interested in this family and have published some revisions and many notes precursory to 
the present treatment. I have to thank the late Mr. N. Y. Sandwith (Kew) for namings of cul- 
tivated species, and Dr. A. L. Gentry (St. Louis) for recent information on them, Dr. H. Heine 
(Paris) for assistance in various matters, Mr. Michael Galore (Lae) and Prof. E. J. H. Corner 
(Cambridge) for photographs, Miss Christine Brighton (Jodrell Lab., Kew) for the first chromo- 
some count in Tecomanthe, while I gratefully acknowledge precursory work performed by Mr. 
J. C. den Hartog on Tecomanthe and Pandorea in 1969/70 at the Rijksherbarium where he 
worked as a graduate student. 

KEY TO THE GENERA 1 

1 . Erect trees or shrubs. 

2. Leaves simple, mostly in whorls 6. Deplanchea 

2. Leaves compound, almost always decussate. 
3. Leaves 1 -pinnate. 
4. Leaf rachis with a sharp, keel-like ridge above. Leaflets 8-12 pairs. Capsules winged 

12. Pajanelia 

4. Leaf rachis not keeled above. Leaflets less than 6 pairs. Capsules not winged. 

5. Corolla salver-shaped, white, nocturnal, the tube 12-18 cm long, narrow-cylindric. Calyx 

spathaceous to the base, 3-6 cm. Leaves nigrescent, with domatia. A flat pseudoseptum developed 

parallel to the valves. Seeds rectangular, with corky wings as thick as the seed . 7. Dolichandrone 

5. Corolla much shorter. No pseudoseptum. Seeds with hyaline wings. Leaves not nigrescent. 

6. Mature leaflets serrate, crenate, or pinnatifid 5. Tecoma 

6. Mature leaflets entire. 
7. Calyx regularly 5-lobed, rather thin, reticulately veined, short-hairy, 2-3 cm 

11. Lamiodendron 1 
7. Calyx lobes unequal, often less than 5; calyx not thin and reticulate- veined. 
8. Capsule rather flattened, with a flat septum, if terete with 10 ribs .... 10. Fernandoa 
8. Capsule terete, with a terete septum, never with many ribs, sometimes one ridge on each 

valve. No domatia in Mai. spp. 
9. Seeds thick, in one row in each cell, each fitting in deep notch in the septum. Valves rather 

hard. Ovules in 2 rows in each cell 8. Stereospermum 

9. Seeds thick, in many rows, the septum without notches. Valves thin. Ovules in many rows 

in each cell 9. Radermachera 

3. Leaves 2-3(-4)-pinnate. 
10. Flowers fleshy, very coarse: calyx 2-4 cm, corolla wide, dirty violet or reddish purple to liver 

brown, 7-10 cm. Capsule flat, sword-shaped, 60-120 cm 3. Oroxylum 

10. Flowers not fleshy, much smaller or thinner, white or pink. Capsule smaller and of different shape. 

11. Leaflets with domatia. Corolla white, salver-shaped, the tube 6-8 cm long, cylindric, 2 mm wide. 

Capsule compressed parallel to the septum, the latter parallel to the valves; dehiscence septicidal. 

4. Millingtonia 
11. Leaflets without domatia. Corolla tube otherwise, wider. Capsule cylindric, with a terete corky 

septum attached perpendicular to the valves ; dehiscence loculicidal 9. Radermachera 

I. Lianas. 
12. Corolla salver-shaped, with a narrow cylindric tube, 5-19 cm long. Capsule flat, large, with a 
median lengthwise ridge. Septum parallel to the valves, dehiscence loculicid .... 1. Nyctocalos 
12. Corolla not salver-shaped, tubular, infundibuliform or funnel-shaped. 

(1) Of Lamiodendron and Hieris the fruit is still unknown. 

The key given here is only to native and thoroughly naturalized species (only Tecoma stans). 
A tentative key to the cultivated species is added in an appendix on page 180. 



1977] Bignoniaceae (van Steenis) 123 

13. Corolla tube — geniculate above the basal tube, upper tube slightly curved and flattened with a 
prominent fold. Calyx c. ■ , cm, below the very short lobes with a short spur-like tooth. Ovules 

6-8 per cell, in — two rows. Leaflets 5, those of the lower pair sessile 2. Hieris 1 

13. Corolla tube not geniculate, without a fold. Ovules cv per cell in several rows. Capsule loculicid, 
with boat-shaped valves. 

14. Leaves digitate, with 3(-5) leaflets. Corolla lobes valvate 13. Neosepicaea 

14. Leaves pinnate, with 1 or more pairs of leaflets. Corolla lobes imbricate. 
15. Flowers in racemes, axillary, mostly on the old wood, rarely terminal (in a high-mountain sp.), 
the rachis at the base usually with several crowded pairs of minute sterile bracts. Calyx large, 
15-40 mm, distinctly lobed. Corolla large, mostly red, 5-12 cm long incl. the lobes, not bearded 
in the mouth and upper part of the tube, but stuppose hairy at the insertion of the stamens (rim 
of the basal tube). Corolla lobes usually deltoid, very narrowly overlapping. Anther-cells 

4-5<-10) mm long 14. Tecomanthe 

15. Flowers in thyrses, only occasionally depauperate into racemes, usually terminal, sometimes 
axillary or on old wood. No sterile bracts at base of peduncle. Calyx small, mostly stunted or 
very shortly lobed, 2-7 mm. Corolla incl. lobes up to c. V , cm (in a single Australian species 
with large lobes c. 5 cm), usually white or yellowish with red dots or streaks inside lobes and 
mouth, often bearded in the mouth and upper part of the tube, sometimes with a hair-ring at the 
insertion of the anthers. Corolla lobes widely overlapping, often roundish. Anther-cells l 1 r 
2 mm long 15. Pandorea 



Tribe 1. Bignonieae 

B. & H. Gen. PI. 2 (1876) 1027; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 209; 
Bureau, Fl. Bras. 8, 2 (1896) 16. — Eubignonieae Endl. Gen. PI. (1839) 712. — 
Subtribe Eubignonieae DC. Rev. Bign., Bibl. Univ. Geneve (1838) 122; Fenzl, 
Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 262; DC. Prod. 9 (1845) 143. 

Capsule septicid, the septum parallel to the valves. Frequently lianas, with 
tendrils, mostly in the neotropics. 

Note. Bojer (Hort. Maur. 218) and DC. (I.e.) included the two present tribes in 
tribe Bignonieae. 

1. NYCTOCALOS 

T. & B. in Miq. J. Bot. Need. 1 (1862) 366; Bureau, Mon. (1864) 52; Miq. Ann. 
Mus. Bot. Lugd.-Bat. 1 (1864) 201 ; ibid. 3 (1867) 249; Seem. J. Bot. 8 (1870) 147; 
K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 219 {'Nycticalos*); Steen. Thesis 
(1927) 805; Bull. Jard. Bot. Btzg III, 10 (1928) 178; Acta Bot. Neerl. 2 (1953) 306; 
Santisuk, Kew Bull. 28 (1973) 182. — Fig. 1-2. 

Lianas, without tendrils. Leaves pinnately 3-foliolate (in one e.xtra-Mal. sp. 
1 -pinnate with 5 leaflets). Leaflets herbaceous to chartaceous, entire, ± elliptic, 
acuminate, with a few scattered crateriform glands along and spaced from the 
midrib underneath, above very fine punctate-pitted glandular. Inflorescence a short 
lateral or terminal 8-12-flowered raceme. Pedicels bracteolate. Flowers nocturnal, 
erect, fragrant; lobes widely imbricate. Calyx cup-shaped, truncate, with 5 horn- 
like teeth, the latter with glands on both sides. Corolla almost actinomorphic, 
saKer-shaped, the long narrow basal tube dilated in the upper part, with 5 rounded 
unequal or subequal lobes. Stamens inserted in the throat 4 with or without a rudi- 
ment or 5, equal or 2 anterior ones sometimes longer, not exserted; anther-cells 
divergent, versatile; filaments glabrous and no hairs near their insertion. Disk 
annular, fleshy. Ovary with x rows of x ovules along the margins of the dissepi- 
ment; stvle long, filiform. Capsule large, flat, stalked and acuminate but with 



124 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 1 . Nyctocalos cuspidata (Bl.) Miq. a. Habit, in flower, b. fruit, opened, showing seeds and dissepiment, 

x »/a (after Miquel, 1867) 

parallel edges, valves with a central prominent rib; dissepiment thinnish, flat. 
Seeds flimsy winged, roundish. 

Distr. Three spp. in SE. Asia (Assam, Burma, Thailand, Yunnan) and West Malesia (Java, Borneo, 
Celebes, and the Philippines). Fig. 3. 

Ecol. Rare rain-forest lianas at low altitude. 

Taxon. The genus stands isolated in the Old World flora. In many aspects the small New World genus 
Tanaeeium Sw. is very similar, but in this genus at least part of the leaflets carries a terminal tendril; 
besides its fruit is not flat, but said to be cylindric or quadrangular with convex woody valves. Still I 
believe it to be an ally in the New World. There is also a similar resemblance with the monotypic 
South American genus Maeranthisiphon Bureau but that has 2-ranked ovules and a more elongate, 
funnel-shaped corolla tube. 

For a moment I thought that Nyctocalos pinnata Steen. (from Yunnan, only known in fruit, I.e. 1953, 
306) might belong to Hieris, but the very numerous seeds defeat this, as far as H. cwtisii is concerned. 
Hieris is, of course, the most intimate related genus, with the same punctate glands on the leaves and a 
deceptively similar calyx; its pollen is quite different. 

Too much importance has been ascribed to the structure of the androecium : 5 equal stamens to 4 
didynamous; this varies as is explained under N. cuspidata. It led Seemann (I.e.) even to the inclusion of 
the Australian Hausmannia jucunda into the genus, which belongs in fact to the Tecomeae with quite 
different fruit and valvate corolla lobes. 

Nomencl. The generic name is female, being derived from the Greek nux. 



KEY TO THE SPECIES 



1. Corolla tube c. 15-16 cm long, white. Calyx teeth horn-like, with a linear apex. Lateral petiolules 
5-12 mm 1. N. cuspidata 

1. Corolla tube c. 5-6 cm long, tinged palish yellow suffused with pinkish shade. Calyx teeth triangular, 
acute. Lateral petiolules 2-3 mm 2. N. brunfelsiiflora 



1977] 



Bignoniaceae (van Steenis) 



125 



1. Nyctocalos cuspidata (Bl.) Miq. Ann. Mus. Bot. 
Lugd.-Bat. 3 (1867) 249, t. 8B {'cuspidatum'); 
Merr. Philip. J. Sc. 1 (1906) Suppl. 237; C. B. Rob. 
ibid. 6(1911) Bot. 211; Merr. En. Philip. 3 (1923) 
443; Steen. Thesis (1927) 813, incl. var. oblongum 
Steen.; Bull. Jard. Bot. Btzg III, 10 (1928) 180; 
Santisuk, Kew Bull. 28 (1973) 183. — Tecoma 
cuspidata Bl. Rumphia 4 (1849) 35. — TV. macro- 
siphon T. & B. Cat. Hort. Bog. (1856) 155, nomen. 
— N. brunfelsiaeflorus (non T. & B.) Miq. Ann. 
Mus. Bot. Lugd.-Bat. 1 (1864) 201, pro specim. 
celeb. — N. thomsonii Hook./. Bot. Mag. 93 (1867) 
t. 5678; Clarke, Fl. Br. Ind. 4 (1884) 377; Steen. 
Thesis (1927) 809; Bull. Jard. Bot. Btzg III, 10 
(1928) 180; Santisuk, Kew Bull. 28 (1973) 183. — 
Gelseminum cuspidatum O. K. Rev. Gen. PI. 2 
(1891) 479. — N. assamica Hook./, ex. K.Sch. in 
E. & P. Nat. Pfl. Fam. 4, 3b (1894) 221, nomen, 
lapsus. — Fig. 1-2. 

Leaflets elliptic, ovate, obovate, or narrow 
oblong, rounded at base, acuminate to cuspidate at 
apex, 6-ll(-18) by 3 1 2 -7(-10)cm; petiole 3-5 cm; 
rachis 2 1 2 -3 1 2 cm; petiolules " 2 -l cm. Pedicels 
c. 1 2 -l cm. Calyx c. 6 mm. Corolla in bud pale 
green, later creamy, the tube c. 15-19 cm; dilated 
part c. 3-6 cm; lobes rounded, c. lVr-2cm. 
Stamens 4, didynamous, with or without a filiform 
rudimentary 5th one; anther-cells 6-10 mm; con- 
nective with a filiform appendage 2-3 mm. 
Capsule 16-24 by 3 3 4 -4 3 4 cm. Seeds (including 
the flimsy wings) rounded to obovate, 3-4 by 
2!-3cm. 




Fig. 2. Nyctocalos cuspidata (Bl.) Miq. Pollen 
grain, SEM • 500 (BS 10396). 



Distr. SE. Asia (Assam: Mikir & Gowhatty 
Hills) and Central Malesla: Philippines (Luzon, 
Polillo, Palawan, Biliran, Mindanao, Basilan) and 
Celebes (Manado, Kema, Bantacng). The type- 
was said to have been collected in the Moluccas by 
Zippm but this must be doubted. Fig. 3. 

Ecol. Lowland rain-Ion 

Notes. Hitherto importance was laid in keys on 
the difference between N. CUSpidatQ and N. 
thnmsomi in that the lonner was described with 5 
fertile stamens and the latter with 4 didynamous 



stamens and a filiform rudiment. This was suspi- 
cious as there were hardly any other differences. In 
material on spirit of Celebes specimens cultivated 
in Hort. Bog. I have now found flowers with 
didynamous stamens whether or not accompanied 
by a staminode. Furthermore, in Riedel s.n. from 
Manado and in BS 10396 from Polillo there are 5 
perfect stamens; but even here in one flower the 
two anterior stamens were somewhat longer than 
the others. There is thus variation in the degree of 
tendency to zygomorphism. This is also visible in 
the difference in size of the corolla lobes of which 
one is mostly larger than the others. The taxono- 
mical implication is the reduction of N. thomsonii. 
In the Malesian specimens the calyx teeth appear 
to be somewhat longer and more horn-like than in 
the Assam specimens depicted by Hooker /. 

2. Nyctocalos brunfelsiiflora T. & B. in Miq. J. 
Bot. Neerl. 1 (1862) 367 (' brunfelsiaeflorus'); Miq. 
Choix (1863) t. vn; Ann. Mus. Bot. Lugd.-Bat. 1 
(1864) 201, excl. syn. et specim. celeb. ; ibid. 3 
(1867) 248, t. 8A; Steen. Thesis (1927) 811; Bull. 
Jard. Bot. Btzg III, 10 (1928) 179 Cbrunfelsiiflorus , )\ 
Acta Bot. Neerl. 2 (1953) 306; Back. & Bakh. 
/. Fl. Java 2 (1965) 536; Santisuk, Kew Bull. 28 
(1973) 183. — N. shanica MacGregor & W. W. 
Smith, Rec. Bot. Surv. India 4 (1911) 280; Steen. 
Thesis (1927) 811 ; Santisuk, Kew Bull. 28 (1973) 
182, 183; Thai For. Bull. Bot. 8 (1974) 88. 

Leaflets elliptic, oblong, or obovate, acuminate 
to cuspidate, 7-13 by 3-6 cm; petiole 4-7 cm; 
rachis l 1 /2~3cm; petiolules 1-2 mm. Racemes 
5-10-flowered. Pedicels \-\ l l 2 cm. Calyx 5-6 mm, 
suffused with reddish tinge in anthesis {ex coll.). 
Corolla whitish afterwards yellowish, later apically 
suffused with pink; tube 5-7 cm, the dilated part 
some 2-2V2cm; lobes rounded to obovate or 
truncate, c. 2 cm. Stamens (so far as known) 5. 
fertile, equal or subequal. Capsule 10-13 by 3-4 cm, 




i [g v Range of the genui Nyctocalos T. & B. : 
\ brunfelslfflora I A B. (triangles), N. cuspidata 

(Hi .) Miq. (dots), and N. ptmata Shin, (circle). 



126 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 4. Hieris curtisii (Ridl.) Steen. a. Habit, X l /n b. corolla, nat. size, c. LS of ovary, disk and recep- 
tacle, x 2V2, d. calyx, x 2 l l 2 , e. CS of ovary, x 7 l / 2 , f. glands on ovary (Henderson s.n.). 



1977] 



Bignoniaceae (van Steenis) 



127 



Distr. SE. Asia (Upper Burma: S. Shan States; 
Thailand: N. & SE.), in Malesia: Borneo (Sabah, 
near Kudat; Sarawak, near Niah), SW. Java 
(Wijnkoops Bay). Fig. 3. 

Ecol. Lowland rain-forests, even in Burma 
below 300 m. 

Uses. Teysmann found it a beautiful ornamen- 
tal; as far as nocturnally flowering plants can be. 
He could easily propagate it by marcottes. Van 
Hasselt (in sehed.) noted that in SW. Java crushed 
leaves are rubbed against head and stomach aches. 

Vern. Kakatjangan, S, SW. Java. 

Notes. N. shanica was distinguished by having 
glabrous anthers; these had in N. brunfelsiiflora 
been described and depicted as hairy to the base. 
This is, however, a lapsus : the base of the filaments 
and tube in the vicinity of the insertion is only 
dotted with small sessile granular glands. Santisuk 



(I.e.) said that the pollen would be different from 
that of N. shanica, but I cannot accept this for 
specific distinction. The colour of N. shanica was 
described as white, but field data enumerated by 
Santisuk mention also creamy white, buds pur- 
plish, and pale yellow flowers. Of N. brunfelsiiflora 
Miquel mentioned them to be pale pinkish 'tirant 
legerement vers le jaune', more purplish to anthesis. 
Backer said: corolla at first white, afterwards 
yellowish; tube apically suffused with red. I do not 
ascribe much importance to these faint colour 
variances, especially nocturnal flowers often dis- 
colour with age. 

In comparing the scanty flowering material of 
both species it seems that in N. shanica the widen- 
ing of the corolla tube starts lower than in N. brun- 
felsiiflora, but I can not accept this for specific 
distinction. 



2. HIERIS 

Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 279, f. 13. — Fig. 4. 

Slender woody twiner. Leaves 1 -pinnate, with (1— )2(— 3) pairs of entire leaflets. 
Racemes (?axillary or) terminal. Pedicels bracteolate. Flowers scattered. Calyx 
cupular, articulated with the pedicel, short- or indistinctly 5-lobed, just below the 
margin with 5 spur-like, upcurved teeth at the base of each lobe, each tooth with a 
few glands on each side of its base. Corolla zygomorphic, the basal narrowed tube 
c. V3 of its length, ± geniculate with the upper 2 / i which is widened, flattened, and 
gently curved; lobes subequal, suborbicular, finely capitate-glandular, papillose- 
hairy outside at base. Stamens 4, didynamous, inserted at the end of the basal tube, 
included, 5th rudimentary; anthers connivent in pairs, divaricate, connective 
apiculate. Disk entire, pulvinate-annular, fleshy. Ovary ovate, subterete, with 2 
grooves, microscopically glandular; ovules in ± two rows of 3-4 in each cell. 

Distr. Monotypic. Malesia: Malaysia: Penang I. (near village on north coast). 

Taxon. Outstanding by the few ovules and the geniculate tube of the corolla. The structure of the 
inflorescence seems to be racemose though flowers are articulated. 

Notes. Unfortunately the fruit and seed of this most interesting plant is unknown; from the cross- 
section of the ovary one might assume it to belong to tribe Bignonieae as the grooves of the ovary corres- 
pond with the edges of the septum, while furthermore the calyx structure is a replica of that of Nyctocalos, 
vsith which genus it seems closest related. In Nyctocalos, however, there is a very large number of ovules in 
each cell, which are very few in the Penang plant, like in the South American genus Memora. By the 
pinnate leaves Uteris shows resemblance with Nyctocalos pinnata Steen. from Yunnan, which is unfor- 
tunately only known in fruit, but has abundant seeds in each cell. 

Mr J. Muller told me (Febr. 1975) that the pollen of Hieris is not in the least resembling the showy 
pattern of Nyctocalos. 



1. Hieris curtisii (Ridl.) Steen. Bull. Jard. Bot. 
Btzg III, 10 (1928) 280, f. 13. Tecoma curtisii 
Ridl. J. As. SOC. Str. Br. 49 (1908) 26. — Pandorea 
curtisii RIDL. Fl. Mai. Pen. 2 (1923) 553, f. 125; 
Sin-.. Nova f.uinca 14 (1927) 301; Thesis (1927) 
846, f. 4(2). Ii«. 4. 

Qbbrota etc, with very many small 

lenticcls, nodes with glands and a dark transversal 
'10 lis 20 cm long; petiole 2' .• 8cm; 
rachis r. 4 5 cm; pctiolulcs of lower pair ol leaflets 
4 12 mm. of upper pair (0 )l 2 mm. Leaflet! 
herbaceous, slightly unequal-sided, ovate-oblong, 
long-acuminate, 5-8 by 2 3cm; nerves 4 5 pairs. 



undersurface with scattered, small, rimmed-cratcri- 
form glands, upper surface with microscopical 
pitted-punctate glands. Rachis rather densely 
flowered, microscopically puberulous, 2-7 cm; 
peduncle 2 cm, with barren bracts. Bracts linear, 
2 mm. Pedicels thin, 5 X mm, with I 2 minute 
bracteolea m the lowei hair. Calyx (■> mm, purplish, 
bluntly S-ribbed, inside with dark red microscopical 

capitate-glandular hairs. Corolla 4'\ 5 cm, lube 
yellow, lobes whitish turning pale lilac, ft I CHI 
2 upper recurved, 3 lower erect; outside (W\ transi- 
tion ol tube and lobes Scattered, rather large, 
brown red glands; basal tube I'/jCm by 2 mm, 



128 Flora Malesiana [ser. I, vol. 8 2 

near the insertion of the stamens scattered micro- Distr. Malesia: Penang I., see above, 

scopical red-tipped, capitate-glandular hairs as in Ecol. Not well noted, 3 collections, all from 
the calyx tube. Filaments glabrous, 12 and 14 mm; 1898-1902; fl. June, July, Nov., and a cultivated 
rudiment 5 mm, with reflexed apex; anther-cells specimen in Hort. Sing., Lawn 0, in Febr. 
2V2 mm, connective appendage linear, 1 mm. 
Ovary elliptic l'/z by 1 mm; style 2 cm; stigmatic 
lobes elliptic, IV2 mm- 

3. OROXYLUM 

Vent. Dec. Gen. Nov. (1808) 8; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 225 
('Oroxylon'), I.e. 212 in clavi; Steen. Thesis (1927) 816; Bull. Jard. Bot. Btzg III, 10 
(1928) 181. — Calosanthes Bl. Bijdr. (1826) 760; DC. Prod. 9 (1845) 177; Bureau, 
Mon. (1864) 45, t. 9. — Hippoxylon Rafin. Sylv. Tellur. (1838) 78, nom. Meg. — 
Fig. 5, 7. 

Glabrous tree, robust in all its parts. Leaves 2-3(-4)-pinnate, all nodes with in 
sicco shrinking articulations; leaflets entire. Flowers very large, fetid, nocturnal, in 
large terminal racemes (by exception in a thyrse). Calyx persistent, not articulated, 
coriaceous, closed in bud, with a fine apical pore, later opening campanulate, ± 
entire. Corolla funnel-shaped, lobes 5, subequal, imbricate in bud. Stamens 5, 
subequal, all fertile; anthers 2-celled, cells free, ± parallel. Ovary with cv> rows of 
ovules in both cells. Capsule flat, very large, sword-shaped, linear; dissepiment 
flat, coriaceous. Seeds large in oo rows; insertion linear, 1 cm wide. 

Distr. Probably monotypic. From Ceylon, the Deccan and Himalayas through SE. Asia (also in S. 
China: Yunnan, Kwangsi, Setchuan, Kweichou) and Malesia eastwards to the Philippines, Celebes, and 
Timor. Fig. 6. 

Ecol. A characteristic, short-lived nomad tree, nowhere gregarious, not in mature rain-forest but always 
in openings, secondary growths and thickets, rather indifferent to climate (also in teak forest under 
seasonal conditions) and soils, mostly below 1000 m, but in S.China up to 1375 m (Handel-Mazzetti). 

Taxon. A second species has been described, raised from seed, collected by A. Henry in Yunnan, in 
1889, in the Arnold Arboretum, and named O. flavum Rehder (in Sargent, Trees & Shrubs 1, 1904, 
193, t. 92). Rehder discriminated this from O. indicum chiefly by the sulphur yellow colour of the nearly 
symmetrical flowers, the plain not toothed or crisped corolla lobes, the splitting calyx, and the oblong 
leaflets. 

Several of these characters are not valid, especially if we take into consideration that Rehder's plant 
was an unbranched sapling of 3 m high. In such saplings the leaves are always somewhat longer and 
thinner. The sulphur-yellow corolla is also rarely found in O. indicum from where I described it (1928) as 
var. citrinum Steen. on a cultivated specimen at Bogor so annotated by J. J. Smith (C.H.B. XV.K.B.IX- 
11). The calyx is indeed different from that in O. indicum, in being thinner and having 5 faint ribs, but it is 
lobed by tearing, and this is also sometimes found in fruiting specimens of O. indicum. The corolla in 

0. flavum is also regular and somewhat smaller than usual but an examination of the type showed an 
exactly similar occurrence of hairs at the anther bases, the patelliform glands outside and the granular- 
glandular hairs inside. Remains the plain, entire corolla lobes, and an other character figured by Rehder 
but not mentioned by him, viz that the inflorescence is not a raceme, but a thyrse, with the lower stalks 
5-flowered in double triads and the upper ones in simple triads, a situation never recorded or seen by me 
in O. indicum. I cannot well account for these two differences, but they could be due to cultivation; in 
our experience tropical plants in hothouses often deviate from those in the wild, certainly in first-flowering 
saplings. 

1. Oroxylum indicum (L.) Kurz, Fl. Burma 2 t. 43. — Bignonia indica var. a. Linne, Sp. PI. (1753) 
(1877) 237; Clarke, Fl. Br. Ind. 4 (1884) 378; 625; Roxb. Fl. Ind. ed. Carey 3 (1832) 110. — 
K. & V. Bijdr. Booms. 1 (1894) 66, Atlas 2 (1914) Bignonia pentandra Lour. Fl. Coch. 2 (1790) 379. 
t. 358 ; Ridl. Fl. Mai. Pen. 2 (1923) 548 ; Merr. En. —Bignonia tripinnata Noronha, Verh. Bat. Gen. 5 
Philip. 3 (1923) 444; Steen. Thesis (1927) 816; Bull. (1790) art. 4, p. 8, nomen. — Spathodea indica 
Jard. Bot. Btzg III, 10 (1928) 181, mc/. var. citrinum Pers. Syn. 2 (1807) 273. — Calosanthes indica Bl. 
Steen. I.e. 184; Ochse &Bakh. Ind. Groent. (1931) Bijdr. (1826) 760; Wight, Ic. PI. 4 (1850) t. 1337- 
77, f. 46; Hand.-Mazz. Symb. Sin. 7 (1936) 888; 1338; Miq. Fl. Ind. Bat. 2 (1858) 752; Bureau, 
Corner, Ways. Trees (1940) 166, Atlas t. 29. — Mon. (1864) 45, t. 9. — Bignonia quadripinnata 
Palega-pajaneli Rheede, Hort. Mai. 1 (1686) 77, Blanco, Fl. Filip. (1837) 499, ed. 3, t. 219. — 



1977] 



Bignoniaceae (van Steenis) 



129 




Fig. 5. Oroxylum indicum (L.) Kurz, the 'midnight horror'. Pole on left is a sapling that has flowered 

and fruited and is temporarily leafless. Different branches of the same tree may be in leaf, flower or fruit 

at the same time (Tg. Bukit, Sg. Sedili Ketchil, photogr. Corner, June 1934). 



Hippoxylon indicum Rafin. Sylv. Tellur. (1838) 
78, nom. Meg. — Arthrophyllum ceylanicum Miq. 
Ann. Mus. Bot. Lugd.-Bat. 1 (1863) 27. — Arthro- 
phyllum reticulation Bl. ex Miq. I.e., et corr. 318. — 
l-i«. 5, 7. 

Smallish, glabrous, sparingly branched, semi- 
deciduous tree, 6 20( 27) m; trunk 10-40cm 0, 
with grey bark and large leaf-scars; twigs thick, (as 
the trunk at least at apex) pithy, later hollow, 
Icnticcllate, as the leaf-rachis. Leaves tufted at 
twig-ends, with a long petiole, '/* 2 m; leaflets long 
peddled, ovate to oblong, acuminate, 4 IK 15) by 
i 9cm, cuneaie, rounded or reniform at the trip- 
lincrvcd mostly oblique base, underneath distinctly 



reticulate-veined, with some scattered gland fields 
near the axils of the nerves and scattered, micros- 
copical scales. Innovations of leaves and racemes 
viscid. Racemes terminal, erect, */ 4 — 1 "/a m long, pith 
of twig-apex, peduncle and rachis partitioned. 
Pedicels long, with a few bracteolcs in lower part, 
2-4 cm. Calyx coriaceous, becoming almost woody 
in fruit, containing water in bud, truncate or 
irregularly shallow lobed by tearing, campanulas, 
brown or dirty-violet, 2-4 by l'/2-2cm. Corolla 
reddish purple to liver-brown to dirty violet out- 
side, dirty yellowish to pinkish inside, with a foxy 
stench, 7 10 cm long, the lobes subequal, in young 
buds strongly folded into a massive apex, I 



130 



Flora Malesiana 



[ser. I, vol. 8 2 



crisped or undulate-crenate, in flower patent to ± 
reflexed, outside with scattered patelliform glands, 
the lobes inside with dense, almost sessile capitate 
gland-hairs; basal tube wide, widened to base, 
c. l l / 2 cm. Stamens inserted in throat, their base 
long hairy. Style 4-6 cm, dark violet as the sub- 
entire, large disk. Capsule pendent, 45-120 by 
6-10 cm, valves flat, almost woody, finally black. 
Seeds incl. wings 5-9 by 2Vz-4 cm. 
Distr. As the genus. Fig. 6. 




Fig. 6. Range of the genus Oroxylum Vent. 



Ecol. As the genus. As a consequence of its 
short-lived, short-sized nomad habit relatively rare 
in tracts with largely high primary forest, e.g. 
Borneo. In Malaya chiefly by villages and by rice- 
fields (Corner). Also not particularly common in 
open but seasonally very dry tracts, such as the 
Lesser Sunda Is. and in teak forest largely confined 
to mixed forest stands. Fl. Jan.-Dec, according to 
Koorders in Java at the start of the dry season ; 
fr. July-May, the fruit remaining during the dry 
season on often leafless stems. 

Corner (I.e.) gave a lively account of his obser- 
vations on this grotesque treelet. He remarked: 
"that each leaf develops as a unit and when it 
withers it breaks up gradually in regular order from 
the tip to the base: the leaflets fall off singly and 
the main stalk and its side-stalks break up at the 
joints: the bits accumulate round the base of the 
trunk like a collection of limb-bones, so that we 
may call it the 'Broken Bones Plant'. The leaves 
are crowded near the end of the stem or its 
branches, and saplings, which remain unbranched 
until after their first flowering at a height of some 
15 ft., look like gigantic umbrellas. When the 
saplings flower, the inflorescence develops from 
the apical bud and therefore further upward growth 
of the main stem is prevented. When the inflores- 



cence has finished flowering, the leaves below it 
fall off and the leafless stem is left as a pole with a 
few sabre-like pods dangling from its extremity: 
wherefore, we may call it the 'Tree of Damocles'. 
Then, after 3-4 weeks in a leafless state, one or more 
lateral buds on the stem break out and grow into 
side-branches which, in due course, flower, fruit, 
shed their leaves and branch in their turn: and, 
thus, the big trees are constructed sympodially 
with open irregular crown and a few lanky ascend- 
ing limbs. Each branch seems to flower independ- 
ently of the others so that flowers, fruits and grow- 
ing twigs may be found on the same tree." 

Flower biology. The flowers are nocturnal; on 
each raceme 1-2 flowers open on one night. 
According to Corner (I.e.): "The corolla begins 
to open about 10 p.m., when the tumid, wrinkled 
lips part and the harsh odour escapes from them. 
By midnight, the lurid mouth gapes widely and is 
filled with stink. Before sunrise the corolla is 
detached and slips off over the long style. The 
flowers are pollinated by bats which are attracted 
by the smell and, holding to the fleshy corolla 
with the claws on their wings, thrust their noses 
into its throat: scratches, as of bats, can be seen on 
the fallen flowers of the 'Midnight Horror' next 
morning." Fig. 7. 

Dispersal. The gauzy seeds slip out of the opened 
pods and flit away on the breeze with the jerky 
motion of a butterfly : so in noon-tide, we may call 
the tree the 'Midday Marvel' (Corner, I.e.). 

Uses. Popular with the Sundanese as a veget- 
able (lalab), fresh young leaves and flowers; even 
unripe capsule valves are eaten after being cooked 
(Hasskarl). 

In Bawean I. flowers are used against inflam- 
mation of the eyes. The bitter bark is chewed in 
Java for depurative purpose, especially after 
delivery. 

In Sarawak used for dyeing rattan of black shiny 
baskets. 

In West Java (Priangan) local people are con- 
vinced that the tree is a protection of the house 
against thieves, a superstition probably derived 
from the sword-like shape of the capsules. 

Vern. Midnight horror, E; Malaya: (beka) 
kampong, blalai, blonglai (kaya), bulai kaju, kain, 
merlai, poko bulai, Malacca; Sumatra: bolai, 
Minangkabau, habreng, Atjeh, (ka)kapung, M, S. 
Sum., mengleo, Simalur I., abang-abang, Asahan; 
Borneo: gimurai, Sarawak, Bidayan name; Java: 
ki tongtorang, pongpor(r)ang, S, (kayu) lanang, 
mungli, wongli, wungli, J, pedangan, Japara, deleg, 
kadjen djaler, keok, padangan, raon, J (all once 
noted), bunglo, punglo, Md; h. lema kaba, kowa, 
Flores; Celebes: buli, Bantaeng, pohon padang, 
Manado; karu kadang, kayu, Kutai; Philippines: 
balilang-uak, pingka-pingka, p.-pinkdhan, taghilau, 
Tag., abang-abang, P.Bis., abong-abong, Bis., 
sakayan-bakus, Tagb., balay-uak, bunglui, Sul., 
baliuag, bungoi, C.Bis., banloi, Sub., barahgau, 
kamkampilan, Ilk., kampilan, Neg., maidbaid, Bik. 



1977] 



Bignoniaceae (van Steenis) 



131 




Fig. 7. Oroxylum indicum (L.) Kuk/. a. Top of raceme with two open flowers, • '/j. b. bat arriving on a 
flower, c. landed bat 00 I flower sucking honey (Old Hot. Garden, Univ. Malaya, Kuala Lumpur, photogr. 

SotPADMO, Sept. 1973, 9.30 p.m.). 



132 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 8. Millingtonia hortensis L.f. a. Habit, x l / 2 , b. detail of underside of leaflet, showing domatia, x 5,c. 
flower, nat. size, d. anthers, one in CS, e. capsule, x 1 / 2 , f. seed, nat. size (a after Wallich, c-d after 

Bureau, b, ^-ZSpanoghe s.n. Timor). 



1977] 



Bignoniaceae (van Steenis) 



133 



4. MILLINGTONIA 

Linne/. Suppl. (1781) 45, 291, non Donn, 1807, nee Roxb. 1820; K.Sch. in E. & P. 
Nat. Pfl. Fam. 4, 3b (1894) 226, f. 89 j-k; Steen. Thesis (1927) 825; Bull. Jard. Bot. 
Btzg III, 10 (1928) 186. — Nevrilis Rafin. Sylv. Tellur. (1838) 138, nom. Meg. — 
Fig. 8. 

Medium-sized evergreen or deciduous tree with corky bark. Leaves 2-3-pinnate, 
with domatia. Thyrses lax, c\>flowered, terminal. Flowers white, fragrant, noctur- 
nal. Calyx small, truncate-campanulate, ± 5-lobed, persistent. Corolla salver- 
shaped, glabrous, with a very long, slender, basal tube at apex widening towards 
the limb, limb at base short funnel-shaped, zygomorphic (± 2-lipped), 5-lobed, 
imbricate in bud. Stamens 4, didynamous, glabrous, inserted at the base of the 
widened part of the tube (throat), shortly exserted, no staminode; anthers with 
1 fertile cell, the other spur-like, barren; connective dorsal, swollen. Disk cup- 
shaped, crenate. Capsule linear, compressed parallel to the septum, septicid- 
dehiscent, valves flat. Seeds cv>, thinly discoid, winged. 

Distr. Monotypic. SE. Asia (India, Burma, Thailand, Indo-China, Yunnan) and Malesia: probably 
native, in E. Java, Madura and Kangean Is., Lesser Sunda Is. (Bali, Sumbawa, Sumba, Flores, Timor), 
and S. Celebes (SW. Peninsula ; Muna I.), in many places also cultivated (Penang, Sumatra, Java, etc.) and 
in dry areas running wild, may be wild also in N. Malaya (Perlis and Kedah, Corner, I.e.). Fig. 9. 

Ecol. Lowland monsoon forest. 



1. Millingtonia horten >is Linne /. Suppl. (1781) 
291; Decne, Herb. Timor. (1835) 32; Span. 
Linnaea 15 (1841) 326; Miq. Fl. Ind. Bat. 2 (1858) 
753; Bureau, Mon. (1864) 45, t. 8; F.-Vill. Nov. 
App. (1880) 150, cult. Manila; Clarke, Fl. Br. Ind. 
4 (1884) 377; K. & V. Booms. Java 1 (1894) 65; 
Steen. Thesis (1927) 826; Bull. Jard. Bot. Btzg III, 
10 (1928) 187; Corner, Ways. Trees (1940) 165; 
Merr. J. Arn. Arb. 25 (1944) 316; Back. & Bakh. 
/. Fl. Java 2 (1965) 234. — Bignonia suberosa 
Roxb. Cor. PI. 3 (181 1) 1 1, t. 214, nom. Meg. — Big- 
nonia cicutaria Mart. Denkschr. K. Ak. Wiss. 
Munchen 6, Kl. Math. Phys. (1820) 153, t. D. — 
M. dubiosa Span, in Hook. Comp. Bot. Mag. 1 
(1835) 348, nomen. — Nevrilis suberosa Rafin. 
Sylv. Tellur. (1838) 138, nom. Meg. — Fig. 8. 

Evergreen (or deciduous?) tree, 5-25 m, to 
30 cm 0; bark corky, very rough, cracking; twigs 
Ienticellate. Mature leaves herbaceous, nearly 
glabrous, 3-5-jugate, lower pairs pinnate, up to 
1 m; leaflets ovate-lanceolate, acuminate, sinuate 
or crenate, or entire, 2'/2-6 by 1 Va— 3 cm; domatia 
haired. Thyrse erect, 10-^40 cm, puberulous, 
flowers fragrant, only few open at a time. Calyx 
2-4 mm, teeth short, broad, obtuse, margin revo- 
lute. Corolla tube 6-8 cm by 2 mm, widened to 
mouth, limb 4-5 cm 0, lobes ovate, acute, out- 
side with cratcriform glands, c. l'/,cm. filaments 
c. 10 and 14 mm long; anthers 2 mm, with a 
small appendage at the base. Style to 8 cm long. 
Stigmatic lobes ovate-acute, l'/ 2 mm. Capsule 30- 
35 by 1' 2 l 1 4 cm. Seeds thin-discoid, IVj-JVi by 
1-1 */j cm including the wings. 

Distr. As the genus. Fig. 9. 

Ecol. A characteristic tree of regions subject to 
annual drought ('monsoon Bon*), umipanion of 
teak, fire-resistant by its thick corky bark and pro- 




Fig. 9. Range of Millingtonia L. /. ; delimitation in 
Asia is slightly arbitrary. 



fuse capacity of suckering from roots, below 750 m 
alt. In Timor common in Ziziphus stands (Meijer 
Drees). Fl. Jan. -Sept., mostly June. 

Father Schmutz reported that it is in Flores not 
deciduous; flowers appear at the end of the dry 
season, before the first rains set in. 

Uses. The soft, even-grained timber was some- 
times advertized for tea-boxes but is not of high 
quality. Tree sometimes used for parks or road- 
sides, leaves as a poor substitute of opium in 
ug.ueiies, sometimes received from the opium 
factory in Java under the vernacular name gendje. 



134 Flora Malesiana [ser. I, vol. 8 2 

Vern. Indian cork tree, E, kurkboom, D, Notes. From Sumba the flowers have once 

kahombu, M (Sum.), amfiunan, sikar pitak, sikar been noted to be red (Iboet 264), never confirmed. 

putih, J, karpoti, Kangean, kanongoh, Bali, Meijer Drees (Comm. For. Res. Inst. Bogor 33, 

kitangar, Sumba, takah, Dawang lang., Timor, ai 1951, 39) recorded that Millingtonia is deciduous 

katong inggar, takah, toka hau, Timor, ai kakassa, in the driest regions of Timor, but the scant field 

Tetun lang., Port. Timor, katangka, Bug., Makas- notes do not confirm this, 
sar, kaulolo, Muna I. 



Tribe 2. Tecomeae 

Endl. Gen. PI. (1839) 711; Fenzl, Denkschr. K. Bay. Bot. Ges. Regensburg 3 
(1841) 261 ; B. & H. Gen. PI. 2 (1876) 1029; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b 
(1894) 209; Bureau, Fl. Bras. 8, 2 (1897) 300. — Subtribe Catalpeae DC. Rev. 
Bign., Bibl. Univ. Geneve (1838) 123; Prod. 9 (1845) 203. 

Capsule loculicid, the septum attached transverse to the valves. Mostly trees or 
shrubs, rarely lianas, by exception with tendrils. 



5. TECOMA 

Juss. Gen. (1789) 139; Rehder, Mitt. Deut. Dendr. Ges. 22 (1913) 262; Britton, 
Bull. Torr. Bot. Club 42 (1915) 372; Urban in Fedde, Rep. 14 (1916) 313; 
Melchior, Ber. Deut. Bot. Ges. 59 (1941) 18-31. — Stenolobium D.Don, Edinb. 
Phil. J. 9 (1823) 264; Seem. J. Bot. 1 (1863) 87; Steen. Thesis (1927) 964; Bull. 
Jard. Bot. Btzg III, 10 (1928) 217. — Tecomaria Spach, Hist. Nat. Veg. 9 (1840) 
137; Fenzl, Denkschr. K. Bay. Bot. Ges. Regensburg 3 (1841) 266; Seem. J. Bot. 1 
(1863) 19-23; Sprague, Fl. Cap. 4, 2 (1904) 448; Steen. Thesis (1927) 831; Bull. 
Jard. Bot. Btzg III, 10 (1928) 193; Brummitt, Bull. Jard. Bot. Nat. Belg. 44 (1974) 
421. 

Erect or scrambling shrubs or small trees. No gland fields at the nodes. Leaves 
1 -pinnate, sometimes 1 -jugate, or more rarely simple; leaf or leaflets incised or 
serrate, densely microscopically glandular-punctate and with hairy domatia under- 
neath. Pedicel with minute bracteoles. Flowers in terminal racemes or more often 
raceme-like thyrses, yellow, orangish or scarlet. Calyx cupular or campanulate, 
with 5, often apiculate deltoid lobes, glands scattered. Corolla tubular, with a short 
basal tube, funnel-shaped, ± straight or ± curved, widened to the mouth, lobes 
almost equal to unequal, imbricate in bud, minutely ciliate. Stamens 4, didyna- 
mous, exserted or included ; anther-cells divergent, often finally standing out trans- 
versally, free or partly connate, sometimes hairy; 5th rudimentary. Disk cupular- 
pulvinate to shallowly cup-shaped. Ovary narrow cylindric or oblong, compressed, 
lepidote; ovules 2-4-seriate in each cell. Capsule linear, ± compressed parallel to 
septum ; valves smooth. Seeds hyaline-winged all round, insertion punctiform. 

Distr. Some dozen species in the New World, from extreme S. Arizona and S. Florida to northern 
Argentina, especially in the Andes, and one species in southern Africa. Some species widely cultivated in 
the tropics and subtropics and one of these locally naturalized in Malesia. 

Tax on. I can see not sufficient reason to keep Tecomaria generically apart from Tecoma. It often is said 
to differ by the exserted stamens and orange-red to scarlet flowers, but it has appeared that among the 
many taxa of South American Tecoma (Stenolobium) there are taxa with exserted stamens and in some 
the flowers are orangish. Inadvertently Seemann (J. Bot. 1, 1863, 19-23) also united them, but curiously 
later distinguished Stenolobium (I.e. 87). 



1977] 



Bignoniaceae (van Steenis) 



135 



According to Sprague (Fl. Cap. 4, 2, 1904, 448) there are only two valid characters, viz the number of 
the rows of ovules in each cell (2 in Tecoma, 4 in Tecomaria) and the anthers. As to the first character, in a 
dozen American genera this number varies, from 2-A, 2-6 and in Tabebuia even from 2-many; its 
value seems therefore to be rather low. The second character holds : in American Tecoma the anther-cells 
are completely free causing them in full anthesis to stand often perpendicular to the filament ; in Tecomaria 
they are connate in the upper 3rd or 4th part, so that they can not diverge so widely. 

The intimate relationship between Tecoma and Tecomaria is emphasized by a reputed fertile hybrid, 
x Tecoma smithii W. Watson (see p. 118), between Tecoma velutina and Tecomaria capensis. 

As the differences between the genera coincide with the geographical disjunction I am prepared to 
distinguish them at sectional level and refer Tecomaria to Tecoma sect. Tecomaria (Spach) Endl. Gen. 
PL (1839) 71. 

From southern Africa 3 spp. were described but F. White (For. Fl. N. Rhod. 1962, 380) and Brummitt 
(Bull. Jard. Bot. Nat. Belg. 44, 1974, 419) distinguish only one. 

In South America a thorough revision probably will also show reduction to fewer variable and raciated 
species. 



1. Tecoma stans (L.) H.B.K. Nov. Gen. Sp. 3 
(1819) 144; DC. Prod. 9 (1845) 224; F.-Vill. Nov 
App. (1880) 151; Merr. FL Manila (1912) 428 
Johnston, Proc. Cal. Ac. Sc. IV, 12, 2 (1924) 1 166 
Sandwith in Pulle, Fl. Surinam 4, 2 (1938) 
79; Corner, Ways. Trees (1940) 170, f. 44, 
pi. 159; Back. & Bakh. /. FL Java 2 (1965) 539; 
Gentry, Ann. Mo. Bot. Gard. 60 (1973) 958, 
f. 38, with full synonymy. — Bignonia stans 
Linne, Sp. PL ed. 2 (1763) 871 ; Juss. Gen. (1789) 
139; Rechinger, Denkschr. K. Ak. Wiss. Wien 85 
(1911) 356. — Stenolobium stans Seem. Ann. Mag. 
Nat. Hist. 10 (1862) 30; J. Bot. 1 (1863) 88, incl. 
var. pinnata Seem, type var.; Bureau, Mon. 
(1864) t. 13; Merr. En. Philip. 3 (1923) 444; Steen. 
Thesis (1927) 905; Bull. Jard. Bot. Btzg III, 10 
(1928)218. 

Shrub, up to c. 1-4 m. Leaflets 1-3 pairs (cult, 
sometimes 1-foliolate) lanceolate, acuminate, 
serrate, glabrous, but often along midrib laxly 
hairy, 3-10 by 1-4 cm, cuneate at the base, no 
proper petiolules; petiole 2-5 cm. Racemes 
glabrous, c. 5-15 cm. Pedicels 5-10 mm. Calyx 
campanulate, 5-7 mm, usually with some im- 
pressed plate-shaped glands in middle part or 
upper half, lobes short-ciliate. Corolla yellow, 
3 1 2 -5 cm, limb up to 3'/2cm 0. Stamens included, 
anther-cells ± pilose. Capsule acute, often lenti- 
cellate, 10-22 by Vj- 3 /^ cm. Seeds (incl. wings) 2 by 
V, cm, inserted in two rows on the margins of the 
septum. 

Distr. From Florida through Central and South 
America to N. Argentina, widely cultivated in the 
tropics, also in Malesia, and sometimes run wild, 
naturalized e.g. in Tahiti, the Society Is. (Raiatea), 
and the Marquesas (Nunuhiva), often together 
with tree ferns and Gleichenia. 

Notes. Vegetatively a rather variable species. 
The normal form is with pinnate leaves, but some- 
times there are 3-foliolate and even simple leaves 
intermixed in one sheet. In Tahiti the specimens 
have 5-6 pairs of leaflets. (This may be var. multi- 
jugum R. E. Friks, Ark. Bot. 1, 1903, 401). In 
Mexico a sheet had almost entire leaflets (Sumi- 
I iiHAsr 1885). There is in America a form with 



underneath woolly-hairy leaflets: T. stans var. 
velutina DC. (T. mollis H.B.K.), but the density of 
the tomentum varies considerably in degree and I 
am not very much in favour to recognize this ; this 
is also the opinion of Standley (Trees, Shrubs 
Mexico, 1926, 1319). 

Johnston (J. Arn. Arb. 21, 1940, 264) said that 
in Mexico the normal-leaved form occurs in coastal 
regions, the incised-leaved form in inland places and 
the tomentose form south of these two, all three 
replacing, suggesting subspecific segregation. 

It is rather peculiar that, though the normal- 
leaved form is widely cultivated throughout 
Malesia, the only naturalized one is a fairly con- 
stant form with deeply incised leaflets, which 
seems to be rather rare in the Americas. 

var. incisa G. Don, Gen. Syst. 4 (1838) 224; 
J. K. Maheswari, Bull. Bot. Surv. India 3 (1961) 
357. — T. incisa Sweet, Hort. Brit. ed. 1 (1827) 
284, nomen. — T. stans var. apiifolium DC. Prod. 
9 (1845) 224; Back. & Bakh./. Fl. Java 2 (1965) 
539. — Stenolobium stans var. apiifolium Seem. J. 
Bot. 1 (1863) 89; Steen. Thesis (1927) 906; Bull. 
Jard. Bot. Btzg III, 10 (1928) 218. — ? Stenolobium 
incisum Rose & Standley, Contr. U.S. Nat. Herb. 
16 (1913) 174. — ? T. stans var. angustatum 
Rehder, Mitt. Deut. Dendr. Ges. 24 (1915) 227. 
— T. incisa (Rose & Standley) Johnston, J. Arn. 
Arb. 21 (1940) 264. 

Leaflets (2-3-)4(-5) pairs, very coarsely toothed 
to deeply pinnately incised, not rarely to the mid- 
rib, making acute-triangular lobing, 5-10 by 
l-2V2cm (incl. teeth). 

Distr. Central America, widely cultivated in 
the tropics, also in Malesia: naturalized in Timor, 
Ternate, and SW. New Guinea (near Uta); also 
naturalized in the Concan and N. Kanara (Talbot). 

Ecol. In Timor (Kupang and Baucau Plateau) 
characteristic for red calcareous soils and lime- 
stone, in the latter place gregarious in shrubberies, 
flowering already at an early age; 5 500 m. Fl. 
mostly Aug.-Sept. (Oct.),/r. Oct. Dec. 

Vern. Yellow bells, E, Malaya; ai funan, Tetu 
lang., E. Timor; dufa dufa, Ternate. 



6. DKPLANCHEA 

Vn n i ari), Bull. Soc. Bot. Normandic 7 (1862) 96; Bureau, Bull. Soc. Bot. Fr. 9 
(1862) 164; Beauvis. Gen. Montrouz. (1901) 90; Steen. Thesis (1927) 906, f. 8-9; 



136 



Flora Malesiana 



[ser. I, vol. 8 2 



Bull. Jard. Bot. Btzg III, 10 (1928) 218, f. 2-3 ; Heine, Fl. Nouv.-Caled. 7 (1976) 71, 
f. 16-17. — Diplanthera Banks & Sol. ex R.Br. Prod. (1810) 448, non Thouars, 
1806, nee Schrank, 1819; Scheffer, Nat. Tijd. N. I. 31 (1870) 332; B. & H. Gen. 
PI. 2 (1876) 1048; Steen. Nova Guinea 14 (1927) 293. — Bulweria F.v.M. Fragm. 
4 (1864) 147. — Fig. 10, 12-13. 




Fig. 10. Deplanchea bancana (Scheffer) Steen. a. Flower, b. fruit on thickened rachis, c. opened capsule, 
seeds covering the septum, d. seed, all nat. size. — D. glabra (Steen.) Steen. e. Flower, nat. size, f. LS of 
ovary, enlarged. — D. tetraphylla (R.Br.) F.v.M. g. Flower, nat. size(a-^C.H.B.XIII-J-65,e-/GjELLERUP 

583, g after Van Steenis, 1927). 



Trees, with thick, pithy branches, robust in all parts. Leaves simple, in whorls of 
3~A, ± tufted to end of branches, entire, at base above with a few large crateriform 
or saucer-shaped glands, underneath often fine-punctiform dotted and sometimes 
with scattered larger crateriform glands; glabrous or with a yellow indument of 
simple hairs. Thyrses terminal, erect, a thick short rachis with crowded, horizontal, 
long-stalked triads or twice-forked triads. Flowers erect, showy, yellow, stalked. 
Calyx on a 2-3 mm high, obconical, solid hypanthium, articulate with the pedicel, 
closed in bud, with short lobes hairy at tip, in anthesis with 5 rather regular, acute 
lobes or tearing into 2-5 unequal, irregular lobes, inside fine-glandular, outside 
not rarely with few large crateriform glands, tip penicellate. Corolla imbricate in 
bud, lobes ciliate, zygomorphous to degree, hardly with a distinct basal tube, tube 
just or far exceeding the calyx. Stamens 4, didynamous, exserted, rarely a 5th 
rudiment, ± erect or recurved to one side, together with a style inserted shortly 



1977] Bignoniaceae (van Steenis) 137 

above the base of tube, base capitate-glandular hairy; filaments ribbon-shaped; 
anther-cells free, wide-divergent. Disk annular, crenate. Ovary subsessile, glabrous, 
2-celled, each cell with 2 closely placed placentas; style very long; stigma with 2 
narrow lobes. Ovules cv>, in many rows. Capsule short-stalked, ellipsoid, with hard, 
boat-shaped valves, erect; septum flattened, lens-shaped, thick. Seeds very many, 
roundish, very thin hyaline-winged all around, punctate-inserted. 

Distr. Probably 5 spp., 1 in West Malesia, 2 in New Guinea (of which 1 sp. also in N. Australia and the 
other also in E. Borneo and Central Celebes), and 2 in New Caledonia. Fig. 1 1 . 

Ecol. Rain-forests with preference for light and secondary forest, kerangas forest, others in woodland 
savannahs and invading grasslands, from sea-level to 1000 m. 

Notes. The much increased collections gave a better understanding in specific delimitation and varia- 
bility of characters, leading to reduction in the number of species. Especially the hairiness occurs to degree 
and is occasionally deviating; in occasional specimens of D. bancana the calyx may possess dense long 
hairs inside the calyx. For this reason I have reduced D. tubulosa Steen. and D. coriacea Steen. The 
Australian D. hirsuta Bailey I have reduced tentatively to D. tetraphylla; I believe it to be a juvenile form 
which accounts for its sinuate leaf margin and occurrence of deviating phyllotaxis, decussate or whorls of 3. 

For brevity's sake the characters mentioned in the key are not repeated in the descriptions. 

Specimens in fruit or in bud, or without corolla are difficult to identify. 

Affinity. Deplanchea has no affinity to other Old World genera. Bureau (Mon. 1864, 51) compared 
it with the genus Delostoma from Andine South America with which it shares several characteristic 
features: thick twigs, terminal inflorescences, simple leaves, and boat-shaped fruit valves. Delostoma 
differs in having the valves said to be unequal, one flat, one boat-shaped, and further by triplinerved leaves, 
a regular, dentate (sometimes 'double') stunted calyx, and pink or violet flowers. 



KEY TO THE SPECIES 

1 . Corolla tubular, the tube ± twice as long as the calyx, straight or slightly curved. Stamens and style 

erect, ± straight in anthesis. Leaves in whorls of 3, underneath almost always very laxly hairy on 

midrib and nerves as is the petiole. Calyx lobes ± equal, corolla lobes ditto .... 3. D. glabra 

1. Corolla tube only for 2-5 mm exceeding the calyx, the limb distinctly zygomorphous, 2 lobes higher 

connate, patent and longer than the others. Stamens and style patent-curved over this lobe or recurved. 

Leaves in whorls of 3-4. 

2. Branches of the thyrse triads, or flowers solitary. Pedicels 1 ' \-2 l U cm long. Bud ± cylindric in shape, 

rarely pear-shaped, often with 5 faint ribs below the lobes. Calyx with a few to several large crateri- 

form glands, the lobes ± equal, in anthesis c. V3-V4 as long as the tube. Corolla tube almost cylindric, 

c. 10 by 5 mm. Stamens c. 3-3'/ 2 cm. Leaves in whorls of 3 (by exception 4), hairy underneath or 

glabrous 1. D. bancana 

2. Branches of the inflorescence often 2(-3) times forked. Pedicels V 2 -2 cm. Buds pear- or spindle-shaped, 
or obovoid, smooth. Calyx without crateriform glands, distinctly widened ± campanulate in anthesis, 
the lobes mostly unequally tearing, sometimes only 2 or 3, 1 l 2 - 2 l3 as long as the tube in anthesis. 
Corolla tube widened almost from the base, c. l-l 1 /* cm h »g h - IV2 cm wide at the mouth in anthesis. 
Stamens 4-4 1 2 cm. Leaves in whorls of (3-)4, always hairy underneath .... 2. D. tetraphylla 

1. Deplanchea bancana (Scheffer) Steen. Thesis orange, darker than corolla. Fruit 10-14 by V/ 2 cm. 

(1927) 921, incl. var. glabra Steen. I.e. 923; Bull. Seeds c. 3 / 4 cm 0, incl. wings 3 by 2 cm. 

Jard. Bot. Btzg III, 10 (1928) 221, f. 2b, 3. — Distr. West Malesia: Sumatra (Palembang. 

Diplanthera bancana SCHEFFER, Nat. Tijd. N. I. 31 Asahan, Bencoolen, Indragiri, Tapanuli), Riouw 

(1870) 334; Hassk. Flora 53 (1870) 219; Clarke, Is. (Karimon I., P. Temiang, P. Kedondong), 

Fl. Br. Ind. 4 (1884) 385; Ridl. Fl. Mai. Pen. 2 Malaya (also Penang I.), Banka (common), 

(1923) 552. — D. coriacea Shin. Bull. Jard. Bot. Billiton, and Borneo. Fig. II. 

Btzg III, 10 (1928) 224, f. 2c e. Fig. lOa-d. Ecol. In primary and secondary forests, in 

Small to large tree, 4 36 m; bole to 20 m; d.b.h. Borneo not rare in heath forest, mostly on sandy 

15 150cm, with small or larger buttresses; bark soils, podsols and wet kerangas, slopes of podsol 

finely fissured, flaky, wood "It. unite. Leaves terraces, from sea-level up to 1000 m. Fl. Jan.- Oct. 

chartaceous to coriaceous, obovate to elliptic, 9-34 V'crn. Mindjanbtng, mtngkuoing, mingkubung, 

by 5 1 'j 20 cm, apex rounded, rarely short-wide- M (Banka), kqyu chindiru, Malacca, labu, Palem- 

acuminatc; base cuncate to cordate; glabrous to bang, mingkubong, mCrtapa, P. Temiang, kayu ri 

yellow hairy in various degree, as is the thyrse; martini, halm, Matak, tui, M (P. Karimon. in error 

petiole 3 6 cm. Peduncle 5 20 cm; rachis 2 5 cm; with Dolicliumlinnr'), indjabletlgien, Billilon. 

primary lateral stalks ..I triad* 2 Jem ( <il\\ 12 Notes A lauK variable plant. In addition to 

IK nam. Corolla tube inside at bate densely capitate* the yeUow-tomentcse 01 velutinoua haired typical 

glandular hairy on insertion ol Anther* form as described by Sou hi k there occur glab- 



138 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 11. Range of the genus Deplanchea Vieillard: 
D. tetraphylla (R.Br.) F.v.M. (dots), D. glabra 
(Steen.) Steen. (triangles), D. bancana (Scheffer) 
Steen. (line). In New Caledonia 2 endemic species. 

rous forms, but in degree, with few hairs, or the 
hairs still more reduced and confined to the lower 
part of the petiolar groove, or only to the axillary 
bud. There are also specimens of which the calyx is 
inside long pubescent with 1 -seriate hairs, notably 
S 11989, 16427, 17591, 25411, SAN 32195, 
Anderson 8398, Hallier B 2507, van Niel 4019, 
but they are hairy as the type or glabrous, or with 
small glabrous leaves, rounded leaves or with short 
acuminate tip. This is also the reason that I cannot 
maintain D. coriacea, as the leaf-base varies from 
cordate to rounded to cuneate, without correlation 
with other sets of characters. Also the number of 
crateriform glands varies and these are also found 
in other specimens. I will not exclude the possibility 
that in the field certain biotypes may be bound to 
certain soil types, but I see no possibility to defini- 
tions and formal recognition from the herbarium. 

In Anderson 8398 from Sarawak the leaves are 
in a whorl of 4. 

Fruits are extremely scarce in the herbarium. 

2. Deplanchea tetraphylla (R.Br.) F.v.M. Second 
Syst. Cens. Austr. PI. 1 (1889) 167; Steen. Thesis 
(1927) 916, incl. var. novoguineensis Steen. I.e. 917; 
Bull. Jard. Bot. Btzg III, 10 (1928) 220; Proc. R. 
Soc. Queensl. 41 (1929) 55; Webbia 8 (1952) 435. 
— Diplanthera tetraphylla R.Br. Prod. (1810) 449; 
Bth. Fl. Austr. 4 (1869) 540; Scheffer, Nat. Tijd. 
N. I. 31 (1870) 335; Banks & Sol. 111. Cook's Voy. 
2 (1901) 72, t. 229; Bailey, Queensl. Fl. (1901) 
1137; Compr. Cat. Q. PI. (1909) 368; White, 
Proc. R. Soc. Queensl. 34 (1922) 52; ibid. 38 (1927) 
259; Lane-Poole, For. Res. (1925) 137; Steen. 
Nova Guinea 14 (1927) 293. — Bulweria nobilis- 
sima F.v.M. Fragm. 4 (1864) 147. — D. bulwerii 
F.v.M. ibid. 5 (1865) 72, (1866) 214. — ? Diplan- 
thera hirsuta F. M. Bailey, Bot. Bull. Dep. Agric. 
Queensl. 14 (1896) 11; Queensl. Fl. (1901) 1137; 
Compr. Cat. Q. PI. (1909) 368. — Faradaya 
chrysoclada K.Sch. & Laut. Nachtr. Fl. Schutzgeb. 
(1905) 370; Beer & H. J. Lam, Blumea 2 (1936) 
225, cf. Lam & Meeuse, Blumea 3 (1938) 201. — 
D. hirsuta (F. M. Bailey) Steen. Proc. R. Soc. 
Queensl. 41 (1929) 56. — Fig. lOg, 12-13. 

Tree, without buttresses, 4-25 m; d.b.h. 10 to 
over 100 cm; bole 1-17 m; bark grey or grey- 
brown, corky, furrowed and rectangular-flaking; 
wood pale straw-coloured. Leaves chartaceous to 
coriaceous, usually obovate or oblong-obovate, 



underneath yellow-velutinous, base somewhat 
cuneate to stunted, exceptionally cordate, on the 
base above with 1-7 cup-shaped large glands, 
ll-23(-60) by 7-14(-30) cm; petiole 2V 2 -5 cm. 
Peduncle 4—12 cm; rachis 3-9 cm; branches 2-7 l / 2 
cm; pedicels 1-2 cm. Calyx 12-14 mm. Fruit 5-11 
by c. 2'/ 2 cm. Seeds incl. the wings 2 by l'/jcm. 

Distr. NE. Queensland (incl. Thursday I., 
Fitzroy I.) and East Malesia: New Guinea and the 
Aru Is. (Trangan and Wokam Is.). Fig. 11. 

Ecol. Predominantly in the periodically dry 
belts of New Guinea, also in gallery forest, very 
rarely in rain-forest, almost confined to grassland 
and wooded savannahs and associated with 
Eucalyptus tereticornis (Central Distr.) or Mela- 
leuca, but also in mixed savannahs (Antidesma, 
Schefflera, palms, etc., at Merauke), not rarely 
common, also a pioneer in fired areas, from sea- 
level to c. 600 m, rarely at 1200 m (Mafulu). Fl. 
May-Oct., fr. July-Oct., often flowers and fruits 
together, but fruiting specimens very rare in the 
herbarium. Dwarf specimens may in places flower 
and fruit. 

C. J. Stefels (Verkenningsrapport Berari 
Komebwaller. Mimeo, Fak Fak, 1956, p. 6, 7, 10, 
phot. 2) reported D. tetraphylla from sandy soils, 
often inundated through an impervious subsoil in a 
heathy forest of Melaleuca. 

Field notes: style greenish yellow, filaments 
yellow, anthers brown. According to van Royen 
the flowers have a sourish-sweet scent and are 
eaten by iuries', lorikeets (at Merauke). 

Uses. At Fak Fak the timber is used for prahus 
by the Papuans. 

Vern. Laargola, Trangan, Aru Is., kapul, M, 
Merauke, bas, Sorong, Mooi lang., tembako d'ora, 
Fak Fak, Ersania lang., pwart, Mumuni, Orokaiva 
lang., pakawa, Maipa, Mekeo lang. 

Notes. The phyllotaxis is not constant, several 
specimens have whorls of 3, reminding of D. 
bancana with which this species is closest related. 
In 3 m high saplings (Docters van Leeuwen 
n. 38, cultivated in Hort. Bog. sub XVI.I.F.8) all 
leaves were opposite. 

Similarly saplings in Queensland may have 
opposite leaves and, moreover, narrow oblong to 
lanceolate leaves with wavy, even toothed margin 
(L. S. Smith 12382). Such plants were described 
as D. hirsuta and may precociously flower; also 
very small normal-leaved specimens may flower, 
obviously at an early age, in New Guinea, possibly 
stimulated by open, pyrogenous habitat. Such 
specimens may also sucker. 



3. Deplanchea glabra (Steen.) Steen. Thesis 

(1927) 919, f. 8f, 1; Bull. Jard. Bot. Btzg III, 10 

(1928) 225. — Diplanthera glabra Steen. Nova 
Guinea 14 (1927) 293. — D. tubulosa Steen. 
Thesis (1927) 926, f. 8g, k, m; Bull. Jard. Bot. 
Btzg III, 10 (1928) 226. — Fig. lOe-f. 

Tree, lV 2 -22m; d.b.h. 12-60 cm; bole 3-12 m; 
bark grey, scaly ; mostly ± glabrous in all its parts. 
Leaves obovate-oblong to elliptic-oblong, coria- 
ceous, usually very laxly haired on the midrib (and 
nerves) below, very rarely yellow short-velutinous 
on inflorescence, midrib, nerves and petiole, fine 
dark-dotted beneath, 9-40 by 4'/ 2 -25 cm, rarely 
with some scattered larger glands ; base rounded to 



1977] 



Bignoniaceae (van Steenis) 



139 




Fig. 12. D tp kmekta tetrapkyUa (ILftt.) F.V.M. Inllorcsccnce from above, capsules, partly opened, showing 
dissepiment (II. photogr. Hoogland 4249; fr. photogr. Womi ksi i y, 1956, Sogcri). 



140 



Flora Malesiana 



[ser. I, vol. 8 2 




<t+lu 




%-ifi& 



•*$BK* 



$k%Ofa i 









* ■ « 






1 


y& 








' v * n 










£!3R^SSU 


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Fig. 13. Rather young tree of Deplanchea tetraphylla (R.Br.) F.v.M. in Bot. Garden Lae (photogr. M. 

Galore). 



1977] Bignoniaceae (van Steenis) 141 

cuneate or subcordate, apex rounded; petiole flowering and fruiting already when small, 10-700 

2V2-7 cm. Peduncle 3-7 cm ; rachis 1-11 cm ; triads (-1000) m. 

Vi-2 cm stalked; pedicels 4—15 mm. Bud spindle- Field notes: pedicels red, filaments and style 
shaped to obovoid. Calyx glabrous, very rarely yellowish green, anthers orange, glands on leaf- 
haired on the mid-sepaline ribs, pustular towards base orange. No buttresses. Fl. fr. March-Oct. 
apex but the pustules hardly ever opening as Vern. Celebes: momo, Malili, kalambutoh, 
crateriform glands, 15-17 mm long, lobes 5 sub- Toradja. 

equal, 3-6 by 2 1 2 -5 mm, c. 2 l lr-yii times as Notes. Although this species is usually almost 

short as the tube. Corolla tube inside near the glabrous, except for some lax hairs on the midrib 

stamens capitate-glandular hairy. Placentas in each beneath, Kostermans & Soegeng 444, from Hol- 

cell2, nearly confluent. Capsule 6-9 by c.2-2 l l 2 cm. landia, has very hairy inflorescences, midrib and 

Seeds incl. wings IV2 by 1 cm. petioles, so that obviously the indumentum may 

Distr. Malesia rather common in North New vary as it does in D. bancana. 

Guinea in the vicinity of Hollandia and Mt Also the single Bornean specimen known so far 

Cyclops, also found twice in Central Celebes (Kostermans 21491) is similarly hairy; it was found 

(Malili, Tobela, Palopo) and once in E. Borneo on a mountain ridge at 1000 m alt. 

(Mt Njapa, Kelai R., Berau). Fig. 11. The two specimens from Central Celebes I refer 

It is remarkable that the species is obviously to this species, although one is in fruit and the 

never found in other parts of New Guinea. other in bud, because: the calyx is ± regularly 

Ecol. Both in high forest and in savannah, but lobed, with a few pustules but without crateriform 

more commonly in pyrogenous grassland glands and the small pod and seed do not match 

{Gleichenia-Ischaemum), as a pioneer, often D. bancana; both spp. have leaves in whorls of 3. 



7. DOLICHANDRONE 

(Fenzl) Seem. [Ann. Mag. Nat. Hist. Ill, 10 (1862) 31, nomen; J. Bot. 1 (1863) 226, 
nomen] J. Bot. 8 (1870) 379, nom. cons.; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b 
(1894) 240, f. 92B-D; Sprague, Kew Bull. (1919) 303; Steen. Thesis (1927) 928; 
Bull. Jard. Bot. Btzg III, 10 (1928) 227. — Pongelia Rafin. Sylv. Tellur. (1838) 78, 
nom. rejic. — Dolichandra sect. Dolichandrone Fenzl, Denkschr. Bay. Bot. Ges. 
Regensb. 3 (1841) 265. — Fig. 15-16. 

Trees with 1 -pinnate leaves (or scattered simple leaves, extra-Mai.), leaflets 
entire (or serrulate, extra-Mai.). Flowers in few-flowered terminal racemes, salver- 
shaped, white, fragrant, nocturnal. Calyx not articulate, closed in bud, later 
spathaceous, caducous. Basal tube of corolla long, narrow-cylindric, upper part 
inflated, lobes mostly crisped. Stamens 4, didynamous, 5th rudimentary, inserted at 
the throat; anther-cells divergent. Disk annular. Ovules oo in 4-6 rows, inserted 
on 2 placentas in each cell, on the septum. Capsule elongate subcylindric to ± 
compressed, septum very narrow, false septum very broad and parallel with the 
valves. Seeds hyaline-winged, in the Mai. sp. rectangular with thick corky wings. 

Distr. Spp. 9, one in tropical E. Africa, 4 in tropical SE. Asia, 3 in tropical N. Australia, and one 
ranging widely from Malabar through Indo-Malesia to New Caledonia. Fig. 14. 

Ecol. All inland species, except the wide-ranging D. spathacea which is a back-mangrove tree. 

Typif. In the Code and in Ind. Gen. D. spathacea (L.f.) K.Sch. has been accepted as the type, following 
Seemann (J. Bot. I, 1863, 226). However, the lectotype must be chosen from the original materials 
incorporated bj I BMZI m Dolichandra sect. Dolichandrone, elevated by Seemann to generic rank. He 
referred to Spathodea b of I NDI U hi k and Spalhodea R.Br. Under the first reference D. spathacea is not 
represented, at most a Dolichandrone represented by Bignoniu spathacea (non L.) sens. Roxb. Corom. 
t. 144, a wrong identification for D. falcate (Wall, ex DC.) Seem. (cf. Sprague, Kew Bull. 1919, 308). 
This then must be the type of the genus. 

• n. The Australian species arc- in habit very different from the African and Indo-Malcsian ones: 
characteristic shrubs or sin. ill trees of xerophytic habit and xeromorphous structure, the leaves sometimes 
obviously not decussate, coriaceous, sometimes entire, with fine parallel ascending veins, or leaflets even 
nccdlc-likc. In addition I find the pods terete and the pseudoseptum not Hat, but irregularly, corky swollen 
with deep impressions Of the seeds. According to I FRBAN (Her. I)cul. Bot, Ges. 34, 1916, 755) these species 
lid also be different in pollen from the other species. I have arranged t hern in subg. Coiinccnc Shin. 

| rhesb, 1927, 931, f, 10). fhey should probably be better arranged In i separate section, rather than In a 

subgenus. 



142 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 14. Range of the genus Dolichandrone (Fenzl) Seem, and its species, the largest range being that of 
D. spathacea (L. /.) K.Sch., a mangrove tree with buoyant seeds. 

Three spp. are distinguished; D. filiformis (DC.) F.v.M. is a fairly constant one, with 3-5 filiform 
leaflets, but D. heterophylla (R.Br.) F.v.M. is very variable, with simple to pinnate leaves (3-7 leaflets) also 
varying in width, whereas D. alternifolia (R.Br.) Seem, with ovate, simple leaflets shows a tendency to split 
the leaf. Already Seemann (J. Bot. 8, 1870, 382) remarked that the latter two probably belong to one 
variable species, which opinion I now tend to share ; he accepted the epithet heterophylla. 



1. Dolichandrone spathacea (L./.) K.Sch. Fl. Kais. 
Wilh. Land (1889) 123; Merr. Fl. Manila (1912) 
429; Int. Herb. Amb. (1917) 469; Sp. Blanc. (1918) 
349; Sprague, Kew Bull. (1919) 304; Steen. 
Thesis (1927) 937; Bull. Jard. Bot. Btzg III, 10 
(1928) 227; C. T. White, J. Arn. Arb. 10 (1929) 
265; Merr. Comm. Lour. (1935) 355; Corner, 
Ways. Trees (1940) 163, Atlas pi. 26-27; Heine, 
Fl. Nouv.-Caled. 7 (1976) 81, pi. 18. — Niir 
Pongelion Rheede, Hort. Mai. 6 (1686) 53, t. 29. — 
Lignum equinum Rumph. Herb. Amb. 3 (1750) 73, 
t. 46. — Bignonia spathacea L.f Suppl. (1781) 283 ; 
Retz. Obs. Bot. 5 (1788) 5; Blanco, Fl. Filip. 

(1837) 499. — Bignonia longissima Lour. Fl. 
Coch. (1790) 380, nom. illeg., non Jacq. 1760. — 
Bignonia javanica Thunb. Mus. Nat. Ac. Upps. 17 
(1794) 150, nomen; Fl. Ceil. (1825) 7, nomen, cf. 
Steen. Blumea 6 (1950) 359. — Spathodea longi- 
flora Vent. Choix (1803) 40; Span. Linnaea 15 
(1841) 326. — Spathodea rheedii Spreng. Syst. 2 
(1825) 835, quoad syn.; Wall. Cat. (1832) n. 6516; 
DC. Prod. 9 (1845) 206; Miq. Fl. Ind. Bat. 2 (1858) 
754. — Pongelia longiflora Rafin. Sylv. Tellur. 

(1838) 79. — Bignonia longiflora Willd. ex DC. 
Prod. 9 (1845) 206. — Spathodea loureiriana DC. 
I.e. 209. — Spathodea luzonica Blanco, Fl. Filip. 
ed. 2 (1845) 350; ed. 3, 2 (1878) 284, t. 242. — 
Spathodea diepenhorstii Miq. Fl. Ind. Bat. 2 (1858) 
754. — D. rheedii Seem. J. Bot. 8 (1870) 380; Kurz, 
Fl. Burma 2 (1877) 234; Clarke, Fl. Br. Ind. 4 
(1884) 379; K. & V. Bijdr. (1894) 69; Gamble, J. 
As. Soc. Beng. 74, ii (1905) 377; Ridl. Fl. Mai. 
Pen. 2 (1923) 549. — D. longissima K.Sch. in E. & 
P. Nat. Pfl. Fam. 4, 3b (1894) 240. — Fig. 15-16. 

For fuller references see Steen. (1928). 



Evergreen, glabrous tree, 5-20 m; 10-40 cm 0; 
wood soft, white. Leaves usually 3-4-jugate, 
15-35 cm, stalked, in the herbarium nigrescent as 
all other parts; young parts ± viscid, young leaves 
slightly pinkish in the field (Corner) ; leaflets thin, 
ovate-oblong to lanceolate, unequal-sided, entire, 
long-tipped (in seedlings sometimes toothed), 6-16 
by 3-7 cm, underneath with hairy domatia. 
Racemes 2-8-flowered. Rachis 2-3 cm. Bracts 
caducous. Bracteoles 0. Pedicels 2-4 cm. Flowers 
not articulated. Calyx conical, coriaceous, usually 
arcuate, beaked, circumscissile caducous, with 
many microscopical glands and a field with large 
crateriform glands at apex, 3-6(-8V2) cm. Corolla 
tube 12-18 cm long, the mouth 7-12 cm ; basal 
tube 9-12 cm, gradually funnel-shaped expanded 
above the throat for 4 cm ; lobes 5, broad, sub- 
equal, frilled round the edge, with large glands, 
2V2-3 cm. Stamens not exserted. Style exserted. 
Capsule flattened-cylindrical, ± ribbed, straight or 
± arcuate, or twisted, tipped, 25-70 by 2-3 cm ; 
valves hard leathery, pseudoseptum flattish, hard 
corky, c. l'/j-P^cm wide. Seeds dark grey, 
rectangular, in many rows, 12-18 by 6-8 mm 
including the thick corky wings ; attachment a fine 
line, 8-10 mm long. 

Distr. From the coast of Malabar throughout 
tropical SE. Asia and the whole of Malesia to New 
Guinea, Micronesia (W. Carolines: Korror; Yap: 
Tomil I.), the Solomons, the New Hebrides and 
New Caledonia, not found in Australia and Poly- 
nesia. Fig. 14. 

Ecol. Confined to the back-mangrove and banks 
of tidal rivers and estuaries. Ridley (Kew Bull. 
1910, 203; J. As. Soc. Str. Br. 59, 1911, 40, 146) 



1977] 



Bignoniaceae (van Steenis) 



143 




Fig. 15. Dolichandrone spathacea (L. /.) K.Sch. Flowers and twigs in bud and fruit, magnification l /, 

upper capsules opened (photogr. Corner). 



recorded it common in low-lying rice-fields near 
Kanga village, Lower Siam, as the predominant 
tree, which he ascribed as "relics of the time when 
this whole country was a tidal swamp, gradually 
filled up after the disappearance of the sea". Other 
seashore plants were also found in these paddy 
fields, such as Euphorbia atoto. This inland occur- 
rence is also stressed by Corner (I.e. 164) who 
found it "frequently in coastal rice-fields; in Perlis 
it is indeed a feature of the country; also in North 
Kcdah, as soon as one reaches Kodiang it attracts 
attention, standing in the paddies as an upright 
poplar and flanks the roads which lead to Kangar 
and Singgora. Old tree trunks are massive and 
fluted at the base, the crown tapering upward. The 
old, opened twisted pods remain for a long time on 
the tree." 



Brass found it very abundant in Daru I. (S. New 
Guinea), while K. J. White found almost pure 
stands in Umboi I. (Morobe Distr.) in swamps 
behind the mangrove. He recorded it also from 
freshwater swamps in the Markham Valley. At the 
Bogor Botanic Gardens it is successfully cultivated 
in freshwater. 

The calyx is filled with water in bud. The very 
young inflorescence and developed ovary is often 
slightly glossy varnished in the herbarium, similarly 
as is found in Radermachera, certainly by the exu- 
date of glands which are found at the apex of the 
calyx. Fl.fr. Jan. -Dec., flowers and fruits not rarely 
found together. Koorders (1894 I.e.) and Heyne 
(Nutt. PI. 1 37 1) say that in the dry season it maybe 
at times nearly leafless fruiting in Central and East 
Java. 



144 



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[ser. I. vol. 8 2 




Fig. 16. Trees of Dolichandrone spathacea (L. /.) K.Sch. in the coastal rice-fields of Perlis where it is a 

feature of the country (photogr. Corner). 



Pollination. Flowers open at dusk and drop 
before sunrise; they must be pollinated by hawk- 
moths with very long tongues to attain the honey. 
In each inflorescence one flower is open at a time 
(Corner). 

Dispersal. The corky seeds float readily and must 
be dispersed by seawater. In this respect it is strange 
that the species is not found in northern Australia 
and Polynesia. 

Uses. Of little use other than fire-wood; in 
N. Borneo a collector deemed the wood useful for 
making clogs and matches ; in the Carolines (Koror 
I.) leaves and fruit are said to be used as a sub- 
stitute for betel leaves in chewing. Heyne (Nutt. PI. 
1371) said that the wood is not durable, but light 
and easy to work for small things in the house; 
pieces of branches are sometimes used for floats of 
fishing nets in East Java and the Karimon Djawa 
Is.; in the Minahassa it is used for scabbards, in 
Madura I. for masks for the toping. In Madura a 
cold concoction of the leaves is also used against 
mouth sprew. Rumphius said that in Ambon twigs 
of lignum equinum (translation of kaju kuda) were 
used for making hedges. 



Vern. Malaya: poko kulo, tuj, M; Sumatra: 
tuwe-ej, Atjeh, kudo-kudo uwi, Simalur, kuda kuda, 
Pariaman, ki arak, Palembang; Java: kaju or ki 
djaran, M, djarang, S, djaram, djaran pelok, 
djaranan, kadjeng kapal, kaju pelok, kapal, J, 
kadju djharan, kaju djaran binek, Md; Borneo: 
kelaju, tuwi, Kutai, toi, tui, Brunei, Bajau lang., 
towi, Kedayan; Celebes: fojet, kaju pelumping, 
sangi, tomana, Minahassa; Talaud Is.: sansarangi; 
Alor: bombila; Ternate: djodjame; Tidore: djame; 
Ambon: kaju kuda, kati kati; Philip.: tua, tui, 
Tag., pata. Ilk., tahgas, Tagb., tanhas, C. Bis., 
tanghas, P.Bis., tewi, Mbo., tiwi, Tag., Bik., C.Bis. ; 
New Guinea: tie, Holtekang, Wembie lang., pide, 
pier, S. New Guinea, Asmat lang., aisumbu, Mani- 
kiang lang., asember tiy, Oransbari, Hatam lang., 
daud, Cape Vogel, Wanigela; New Britain: latiu, 
W. Nakanai, tavituviti, Gazelle Pen., Boava lang.; 
Solomons: kwae kwaele, Guadalcanal, kwe kweale, 
Malaita, kwe'ekwe'eali, Kolombangara, ririge, 
Small Nggela, Kwara'ae lang. 

Note. Especially leaflets of saplings may, at 
times, show some serrulations on the margin and 
may also be slightly hairy. 



1977] 



Bignoniaceae (van Steenis) 



145 



8. STEREOSPERMUM 

Cham. Linnaea 7 (1832) 720; A.DC. Prod. 9 (1845) 210; B. & H. Gen. PL 2 (1875) 
104 pro sect. Eustereospermum ; Steen. Thesis (1927) 946 ; Bull. Jard. Bot. Btzg III, 
10 (1928) 233; Chatterjee, Bull. Bot. Soc. Beng. 2 (1948) 68. — Hieranthes Rafin. 
Sylv Tellur (1838) 79. — Dipterosperma Hassk. Flora 25, 2 (1842) Beibl. 1, p. 28; 
Cat Hort. Bog. (1844) 152; PL Jav. Rar. (1848) 507. — Fig. 17, 19. 




Fi K 17 Stt f o w er nu an ptnonatum (Hawk.) Chattbuto, i HaWt, b. capwle, both l / a , c. teed, nat. 

si/c (waiter Wk.iii. lc. 4, t. 1341. /^ C BBWOEOM A (in sink 3662). 



146 



Flora Malesiana 



[ser. I, vol. 8 2 



Rather tall, deciduous trees. Leaves 1 -pinnate; leaflets a few pairs, diminishing 
in size downwards, underneath usually with flat, dish- or cup-shaped glands or 
glandular spots; no domatia (in Mai.). Thyrses well-branched, paniculiform, mostly 
terminal, or on old wood. Flowers fragrant (in Mai.). Calyx usually short-lobed. 
Corolla infundibuliform ; basal tube mostly concealed in the calyx, upper part 
usually funnel-shaped; mouth bilabiate, upper lip 2-, lower 3-cleft, lobes subequal, 
rounded, crisped, toothed or laciniate. Stamens 4, didynamous, included, 5th 
rudimentary; anthers glabrous, cells divergent. Disk cupular to annular. Ovary 
cells each with 2 rows of many ovules. Capsule long linear, terete, mostly twisted, 
usually 4-angular in section; septum thick, corky, terete, with alternating notches 
to fit the thick seeds which appear in two rows ; valves coriaceous. Seeds co, thick, 
trigonous, wedge-shaped, with a cross-groove, on both sides thinly winged; coty- 
ledons folded, 2-lobed, radicle straight. 

Distr. Over a dozen spp., in tropical Africa and Madagascar, in SE. Asia as far east as Yunnan, in 
Malesia: 2 spp. in Malaya, possibly also in Sumatra, and a doubtfully indigenous record of a third in 
East Java. Fig. 18. 

Ecol. Largely confined to regions subject to a seasonal climate, all in the lowlands, in everwet rain- 
forest obviously deciduous and flowering after a dry spell. 

Note. Besides the 3 Malesian spp. distinguished here, P. Dop mentioned S. cylindricum Pierre from 
Malaya (Fl. Gen. I.-C. 4, 1930, 582), but this must rest on an error as that species is only known from 
Thailand and Indo-China \cf. Santisuk, Thai For. Bull. Bot. 8, 1974, 22). 



KEY TO THE SPECIES 

1. Inflorescence (incl. flowers) viscid-hairy by patent capitate-glandular hairs. Leaves not glabrous. 

Corolla tube gradually funnel-shaped widened, straight, Stamens glabrous at base. Capsule not 

4-ridged. 

2. Leaflets 0-5 mm stalked, base cuneate-attenuate. Capsule faintly 3-ridged on each valve, c. 15-18 mm 

0, septum 8-13 mm 0. Calyx campanulate, 5-7 mm. Corolla dull purple, yellow-streaked within, 

c. 3 cm (stretched); lobes crenulate; tube c. l 3 / 4 cm. Filaments towards the insertion with small 

granular glands 2. S. chelonoides 

2. Leaflets 5-11 mm stalked, rounded to subcordate at the base. Capsule only with 1 median ridge on 
each valve, 8-12 mm 0, septum 6-9 mm 0. Calyx ± cylindric, 8-12 mm. Corolla pale lilac, 6-7 cm; 

lobes deeply fringed-laciniate ; tube c. 4 cm. Filaments glabrous 1. S. fimbriatum 

1. Inflorescence and leaves glabrous (or very rarely minutely puberulous). Corolla c. 3 cm, suddenly 
widened and curved above the basal tube, yellowish, the limb with reddish veins and markings, lobes 
crenulate. Throat and base of filaments densely hairy. Leaflets 5-15 mm stalked, long-acuminate. 
Calyx campanulate, 4-5 mm. Capsule 4-ridged, 8-10 mm ; septum 4-5 mm . 3. S. personatum 




Fig. 18. Range of the genus Stereospermum Cham. 

In Asia and Africa generalized, and numbers of 

species an approximation. Occurrence in Sumatra 

and Java doubtful. 



1. Stereospermum fimbriatum (Wall, ex G. Don) 
A.DC. Prod. 9 (1845) 211; Kurz, Fl. Burma 2 
(1877) 231; Clarke, Fl. Br. Ind. 4 (1884) 383 
Gamble, Man. (1902) 516; J. As. Soc. Beng. 74, ii 
(1905) 378; Ridl. Fl. Mai. Pen. 2 (1923) 550 
Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 234 
Dop, Fl. Gen. I.-C. 4 (1930) 578; Corner, Ways 
Trees (1940) 172, pi. 33; Chatterjee, Bull. Bot 
Soc. Beng. 2 (1948) 69; Santisuk, Thai For. Bull 
Bot. 8 (1974) 23. — Bignonia fimbriata Wall 
[Cat. (1832) n. 6500, nomen] ex G. Don, Gen 
Syst. 4(1838)221. — Fig. 19. 

Very upright, deciduous tree, to 27-30 m by 
30-160 cm 0; crown narrow, cylindrical, rather 
open; bark light grey, rough and flaky; young 
leaves purple or violaceous. Leaves 30-75 cm, with 
rather sticky hairs, stalks yellowish; leaflets 
(2-)3(-4) pairs, rounded at the asymmetric base, 
ovate-oblong, long-tipped, 8-16 by 3-7 cm; 



1977] 



Bignoniaceae (van Steenis) 



147 




Fig. 19. Stereospermum fimbriatum (G.Don) A. DC. 

Its slender habit; some trees deciduous; on the 

churchyard in Malacca (photogr. Corner). 

petiolules 5-10 mm. Flowers in large spreading 
viscid-pilose clusters, I 30 cm 0, on the bare 
twigs or with the new leaves. Calyx tubular, with 
5 very short pointed lobes, c. 8-12 mm. Corolla 
dull white to pale pink or pale pinkish lilac, narrow 
funnel-shaped, without a distinct basal tube, the 
tube 4 5 cm long, the lobes beautifully long-fringed 
(as a dainty night cap), c. 2 cm. Filaments glabrous, 
inserted at r. 11-13 mm from the base CcfSUlt 
more or less quadrangular, snake-like twisted, 
35-60 cm by 8 12 mm; septum r. 5 mm 0. Seeds 



c. 2 l / 2 cm long, 7 mm wide, with rather thick wings. 

Distr. Continental SE. Asia (Burma; Thailand: 
Chiengmai to Peninsular Thailand); in Malesia: 
Malay Peninsula (incl. Langkawi, Penang, and 
Tioman Is.), possibly also in Sumatra. 

The record from Sumatra rests on an un- 
published, beautiful plate in a collection of 
Raffles in the India Office Library & Records 
(NHD 49/20), which might have been made in 
Bencoolen, but according to Mr R. Desmond the 
provenance is uncertain. It might be rare in 
Sumatra, similarly as S. personatum. 

Ecol. In the lowland and hill forests, in Burma 
up to 1000 m, in Malaya in high forest and open 
country: frequent in villages and belukar from 
Malacca to Perlis and Kelantan, often on rocky 
coasts and headlands. Fl. Febr.-June, fr. March- 
Nov. "After the first spell of dry weather the leaves 
are shed and the flowers appear on the bare 
boughs in delicate clusters until the new foliage is 
mature ; in the early morning the corollas spin down 
like snow-flakes and carpet the ground with pale 
lilac blossom. There are many trees in the Christian 
Cemetery at Malacca, and a fairer one for a grave- 
yard would be hard to come by" (Corner, I.e.). 

Uses. A hard and durable fairly large timber 
rather dark coloured, used for beams and posts and 
said to be durable in the soil. 

Roots and leaves are used medicinally for some 
minor ailments: juice of leaves is dropped into the 
ear for ear-ache; leaves pounded with lime are 
applied to the skin for itch ; a decoction of roots is 
given as a protective medicine after childbirth 
(Burkill, Diet. 1935, 2082). 

Vern. Malaya: chac(h)a(h), chachar, chechar, 
chicha(r), M, lempoyan, beka(k) (Burkill), snake 
tree, E (Corner). 

2. Stereospermum chelonoides (L. /.) A.P.DC. 
Bibl. Univ. Geneve II, 17 (1838) 125, pro comb., 
excl. ref. Rheede t. 26; A.DC. Prod. 9 (1845) 210, 
pro basion.; Hmhes, Kew Bull. (1922) 121, in text; 
Santisuk, Kew Bull. 28 (1973) 176; non auct. al. — 
Bignonia chelonoides L. /. Suppl. (1781) 282, pro 
typ., excl. ref. Rheede; non auct. al. — Bignonia 
suaveolens Roxb. Fl. Ind. ed. Carey 3 (1832) 104. — 
Tecoma suaveolens G. Don, Gen. Syst. 4 (1 838) 224. 
— Hieranthes fragrans Rafin. Sylv. Tellur. (1838) 
79 nom. illeg. — S. suaveolens A.DC. Prod. 9 
(1845) 21 1 ; Wight, Ic. 4 (1848) 9, t. 1342; Kurz, 
Fl. Burma 2 (1877) 231; Clarke, Fl. Br. Ind. 4 
(1884) 382; Trimen, Fl. Ceyl. 3 (1895) 284; 
Gamble, Man. (1902) 515; Brandis, Ind. Trees 
(1906) 495; Haines, Fl. Bihar Orissa (1922) 656; 
Beumee, Fl. Anal. Onderz. (1922) 33; Gamble, 
Fl. Madras 2 (1924) 998; Steen. Thesis (1927) 948, 
incl.f. verticillatum STEBN. I.e. 950; Bull. Jard. Bot. 
Btzg III, 10 (1928) 236; Dop, Fl. Gen. I.-C. 4 
(1930) 588; Chatterjek, Bull. Bot. Soc. Beng. 2 
(1948) 70; Back. & Bakh./. Fl. Java 2 (1965) 540. 
Deciduous tree, up to 30 m, 80 cm ; timber 
dark, hard. Innovations viscid hairy. Leavei 
opposite (rarely in whorls of 3), 30 50 by 1 5 25 cm ; 
leaflets 3 4 pairs, viscid-hirsute, glabresccnt, rough 
above and brittle when mature, ovate to obovatc to 
broadly oblong, acute to short-acuminate, entire 

or One-dentate, 3 23 by 3 10 cm: gUtndleM or with 

a few scattered spots; midrib finally puberulous 



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Flora Malesiana 



[ser. I, vol. 8 2 



above, venation sparsely hirsute beneath ; petiolules 
thick, 2-3(-5) cm. Thyrse up to 25 cm 0, viscid- 
pubescent with capitate-glandular hairs. Flowers 
dull crimson to dull purple, yellow streaked within, 
very fragrant. Calyx campanulate, viscid pubescent, 
6-8 mm, shortly acutely 5-lobed. Corolla viscid- 
hairy, 2V 4 -3 cm long, the tube rather gradually 
funnel-shaped, mouth long pubescent; lobes 
subentire, ± as long as the tube. Filaments in- 
serted at 4-5 mm from the base, towards the inser- 
tion with small sessile granular glands. Ovary 
4-ribbed, sometimes sparsely glandular. Capsule 
smooth or valves obscurely 3-ribbed, to 45 by 
l l /2-l 3 /4cm; valves woody; septum 8-13 mm 0. 
Seeds 3V 2 by 3 / 4 cm. 

Distr. Widely distributed in continental tropi- 
cal SE. Asia, from Ceylon and the Deccan to 
Assam and Burma, not yet reported for Thailand 
and erroneously so for Indo-China (Santisuk, 
I.e.); in Malesia: very locally found in East Java, 
but somewhat doubtful whether native. 

Beumee I.e. recorded this tree for the first time 
for East Java, where it was found locally in some 
places in the (teak) forest districts S. Surabaya and 
E. Tuban. He suggested that this occurrence would 
fit the theory of a number of forest officers of early 
import by Hindus of teak and some associated 
trees (Butea monosperma, Schleichera oleosa, etc.) 
and several other plants. I certainly agree that in 
the Hindu period (roughly 800-1400 A.D.) plants 
have come from India, especially those favoured 
for sacred purposes; for example Cochlospermum 
religiosum (L.) Alston, and others went to India 
vice versa, as for example Santalum album L. (see 
C. E. C. Fischer, J. Bomb. Nat. Hist. Soc. 40, 
1938, 458-467). The first is still only found near 
Hindu temples in Bali and the latter is still spread- 
ing in India. The disjunction between the localities 
in East Java and India-Burma is in these cases 
certainly caused by intentional dispersal by man in 
historic time. 

There are, however, a large number of other 
plants showing this same disjunction, and all bound 
to a seasonal climate, that is, subject to a distinct 
annual period of drought. In a succinct analysis I 
found these to belong to 4 classes (Hand. 8th Ned. 
Ind. Natuurwet. Congr. Surabaya 1938, 1939, 
408-409). Later I have further elaborated this 
problem and tried to solve it (Reinwardtia 5, 1961, 
420-429, maps 1-6). From this it appeared that the 
ecological disjunction of the seasonal climate 
between the colossal area it covers in SE. Asia 
(south as far as Tenasserim) and a similar ecology 
in Central & East Java and the Lesser Sunda Is. is 
shared by a homologous plant-geographical dis- 
junction of many hundreds of plants which do not 
occur in everwet West Malesia, or only in very local 
seasonal spots in Celebes and the Philippine 
Islands. A fair number extend their range south- 
eastwards to Australia. This proves that such pat- 
terns are quite natural; I have assumed they 
originated during the Pleistocene Glacial Period, 
which created a temporary pathway for drought 
plants between SE. Asia and Australia, to vanish 
in the Late Holocene. 

It could thus well be that also S. chelonoides does 
occur in the native state in the East Javanese teak 
forest. As a matter of fact no fruit has yet been 
collected, although flowering was abundant. I 



cannot subscribe to the opinion of Beumee that its 
dispersal is here by vegetative means, because I 
cannot well see by what vegetative means and 
furthermore because it is difficult to see how it 
would have maintained itself vegetatively in this 
way for many centuries. On the other hand the 
existence of a Javanese vernacular name is no 
argument that it is native; experience tells us that 
such names are often invented quickly. If it is 
native, it remains curious that, though it is obvious- 
ly of rare occurrence, it was only recently dis- 
covered. It cannot be disproved, however, that its 
seed was inadvertently introduced by the Forest 
Service with teak seed from India or Burma. 

Ecol. Seasonal forest and savannahs. Fl. Sept.- 
Oct. (India: April-June), fr. (Asia) Nov.-Dec. 

Uses. Gamble, I.e. 516, said it is in SE. Asia 
rather an important large tree by its durable timber 
which is easy to work and good for building, 
though the amount of heartwood is small. It also is 
an excellent fire-wood and makes good charcoal. 
The root and bark are used as a favourite tonic 
native medicine. It also is important in sylviculture 
for its very free seed reproduction ; the fruit remains 
long unopened on the tree and seed gets dispersed 
at the very end of the hot season after the danger of 
fire is nearly over, and can germinate with the first 
rains. Even on exposed slopes and among grass its 
good natural reproduction is noticeable. 

Burkill (Diet. 1935, 2082) mentions that it 
yields a gum of the tragacanth class. He also men- 
tioned that it is referred to as a plant of magic in 
Sanskritic India, patala, being the Sanskrit name, 
of which modern vernaculars have been derived. 
In this respect it is noteworthy that the Javanese 
name 'bedalV is a name for Radermachera spp. 
Trimen reported it in Ceylon planted near Buddhist 
temples. 

Vern. Djati teken, kaju teken, J. 

Notes. In East Java one specimen had leaves in 
whorls of three. Leaves of saplings and suckers may 
show a serrate-dentate leaf margin. Such leaves are 
sometimes also rather narrow and acuminate; 
those of mature trees are broader and more wide 
at apex. 

Under S. personatum I referred to the lamentable 
name change caused by erroneous interpretation of 
the type of Bignonia chelonoides L. /. The first to 
observe this was G. Don, I.e., who put B. chelo- 
noides Kon." 1 under the synonyms of Tecoma 
suaveolens. Then Haines remarked that the type of 
Bignonia chelonoides L. /. was currently named 
S. chelonoides (Kew Bull. 1922, 121). I myself 
(Thesis, 1927, 951) was of the same opinion. But 
these observations were not evaluated nomen- 
claturally until recently by Chatterjee and 
Santisuk. 

3. Stereospermum personatum (Hassk.) Chatterjee, 
Bull. Bot. Soc. Beng. 2 (1948) 70; Santisuk, Kew 
Bull. 28 (1973) 178; Thai For. Bull. Bot. 8 (1974) 
26. — Padri Rheede, Hort. Mai. 6 (1736) 47, t. 26. 
— Bignonia chelonoides {non L. /.) Roxb. Fl. Ind. 
ed. Carey 3 (1832) 106, p.p. — S. chelonoides [non 
(L.f.) A.P.DC] A.P.DC. Bibl. Univ. Geneve II, 17 
(1838) 125, quoad ref. Rheede, t. 26; A.DC. Prod. 
9 (1845) 210, p.p.; Kurz, Fl. Burma 2 (1877) 230; 
Clarke, Fl. Br. Ind. 4 (1884) 382; Trimen, Fl. 
Ceyl. 3 (1895) 283; Ridl. Fl. Mai. Pen. 2 (1923) 



1977] 



Bignoniaceae (van Steenis) 



149 



550; Steen. Thesis (1927) 951; Bull. Jard. Bot. 
Btzg III, 10 (1928) 237; Dop. Fl. Gen. I.-C. 4 
(1930) 579; Corner, Ways. Trees (1940) 172, f. 43. 
— Dipterosperma personatum Hassk. Flora 25, 2 
(1842) Beibl. 1, p. 28; Cat. Hort. Bog. (1844) 152; 
PI. Jav. Rar. (1848) 507. — S. tetragonum A. DC. 
Prod. 9 (1845) 210; Haines, Fl. Bihar Orissa (1922) 
655; Kew Bull. (1922) 121 ; Gamble, Fl. Madras 2 
(1924) 998; Hand.-Mazz. Symb. Sin. 7 (1936) 
889. — Bignonia caudata A. DC. Prod. 9 (1845) 
166. — S. hasskarlii Z. & M. ex Zoll. Syst. Verz. 3 
(1855) 54, nom. Meg., based on Dipterosperma 
personatum Hassk.; Miq. Fl. Ind. Bat. 2 (1858) 
756; Ann. Mus. Lugd. Bat. 1 (1864) 200. — 5. 
caudatum Miq. I.e. 200. — Fig. 17. 

Deciduous, glabrous tree, up to 30 m, 75 cm ; 
bark pale pinkish grey becoming rather coarsely 
fissured and flaky but not ridged. Young leaves 
purplish or pinkish (Corner). Leaves glabrous, 
20-50 cm ; leaflets 3-6 pairs, elliptic-oblong, gra- 
dually tapering to the base, tip acuminate to cau- 
date; 5-15 by 2 1 / 2 -6cm; underneath minutely 
glandular-punctate, often with a few large flat 
spot-glands (black in dry state); petiolules slender, 
5-15 mm. Thyrses widely branched, paniculiform, 
to 40 cm long. Flowers dingy yellow, cream within 
with dark red stripes, in slender, erect, lengthening 
thyrses 15-40 cm long, on the bare twigs with the 
new leaves, c. 3 cm long, l 3 4 cm wide at the limb. 
Calyx campanulate, 6-8 mm, with 3-4 short acute 
lobes, purple. Corolla with a narrow basal tube 
4-5 mm long, then campanulately widened and 
curved, the bell-shaped part compressed with the 
mouth closed and the underside grooved; tube 
c. V 4 cm long; lobes crisped, the upper two re- 
curved, the lower bearded at the mouth, ochre- 
buff with brownish or purplish lines, pinkish 
purple on the outside (Corner). Filaments with a 
dense hair tuft at the base. Capsule linear, ± terete, 
4-ribbed, curved or twisted, 8-45 cm by 8-10 mm; 
septum 4-5 mm 0. Seeds 2 by l l 2 cm, incl. wings 
3 « cm. 

Distr. Widely distributed from Ceylon through 
entire continental tropical SE. Asia to Yunnan and 
Indo-China; in Malesia: Malaya (very rare, only in 
Penang, e.g. on Glugor Road, and Singapore), 
possibly also in Sumatra. 

Unfortunately there is no certainty about the 
records in Indonesia: Hasskarl described his type 
from trees cultivated in the Botanic Gardens at 
Bogor; Zollinger described S. hasskarlii also 
from a cultivated tree in these gardens (Zollinger 



3069) but noted that it would have originated from 
Bantam, West Java, adding the Sundanese verna- 
cular l ki langif '; a duplicate of this number in 
Paris is said by Santisuk to have been annotated to 
come from the Lampong Distr. in S. Sumatra, 
which then probably is an error. 

Then there is a collection said to have been col- 
lected by Korthals with 'Borneo' printed labels. 
This provenance is very unlikely, as the use of 
these old labels has been proved to be often erratic. 
These specimens may have come from West 
Central Sumatra but may also have been collected 
by Korthals in the Bogor Botanic Gardens. 
Plant-geographically the species might occur (or 
have occurred) in Sumatra and West Java, but 
probably as rare as in Malaya because of its 
preference for seasonal forest conditions. 

Ecol. Preferring lowland forests, up to 1000 m. 
Fl. March-July (at Bogor Aug., Nov.-Febr.), fr. 
Febr. (Asia), June (Malaya). 

Uses. According to Burkill in India an im- 
portant timber tree, especially in the northeast 
where it is common; the hard grey wood is moder- 
ately durable and easy to work, good for furniture, 
used but less good for building; in Assam and E. 
Bengal padri-wood is used for canoes and tea- 
boxes. In S. India a cooling drink, from roots and 
flowers, is given in fevers. The fragrant flowers are 
offered in temples. 

Nomencl. There has been a most unfortunate 
confusion about the identity of Bignonia chelo- 
noides L. /. (1781). This emanated from Linne /. 
who described it as a hairy plant (type herb. 
Konig, in LINN), but added the reference to 
Padri of Rheede, an other glabrous species with 
long petiolules. This probably misled Roxburgh 
who applied Linne's epithet to the latter. This 
interpretation was followed by almost all subse- 
quent authors. In 1922 Haines concluded that two 
species were involved and he adopted for the 
present one the name S. tetragonum DC. In 1948 
Chatterjee replaced this by an older epithet of 
Hasskarl. 

Notes. As in several other members of this 
family leaves from suckers and saplings may be 
toothed or serrate at the margin ; leaves of mature 
trees are entire. 

In a few continental Asiatic specimens a very 
minute puberulous indument occurs on nerves 
underneath in the inflorescence (Kerr 1167, 
Kostermans 1056, Leschenault 157). 



9. RADERMACHERA 



Zoll. & Mor. in Zoll. Syst. Verz. 3 (1855) 53; Bureau, Adansonia 2 (1861) 192, 
t. 2; Mon. (1864) 50, t. 28; Miq. Ann. Mus. Bot. Lugd. -Bat. 3 (1867) 250; Seem. 
J. Bot. 8 (1870) 145; Jackson, Ind. Kew. 2 (1895) 679 (' RadermachU?)\ Steen. 
Thesis (1927)953; Bull. Jard. Bot. Btzg III, 10(1928) 238; Dop, Fl. Gen. I.-C. 4 
(1930) 583; Chatteveb, Bull. Bot. Soc. Beng. 2 (1948) 71; Santisuk, Thai For. 
Bull. Bot. 8 (1974) 27; Steen. Blumea 23 (1976) 121. — La^aropyxis Miq. Ann. 
Mus. Bot. Lugd. -Bat. 1 (1864) 198. May ode tub; m Klrz, Prel. Rep. lor. Veget. 
Pegu, App. D (1875) pi. 1 & 2; Fl. Burma 2 (1877) 232. — Stereospermum sect. 



150 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 20. Radermachera peninsularis Steen. a. Leaflet, b. apex of thyrse, c. calyx and style, d. corolla, all 
nat. size. — R. glandulosa (Bl.) Miq. e. Leaf base, underneath with dense glandfield, x 2, f. empty 
capsule, with 2 valves and septum, x V2, g- section of septum, x 5, h. seed, x 2 (a-d Larsen c.s. 31239, 

e-h Kostermans 6360A). 



1977] Bignoniaceae (van Steenis) 151 

Radermachera et Xylocarpaea B. & H. Gen. PI. 2 (1876) 1047; Clarke, Fl. Br. Ind. 
4 (1884) 383; K. & V. Bijdr. Booms. 1 (1894) 71. — Radermachera sect. Alatae 
Steen. Acta Bot. Neerl. 2 (1953) 307. — Fig. 20, 22. 

Trees, rarely shrubs, evergreen (except 2 Indian and 2 Chinese spp.). Twigs and 
floral rachis mostly distinctly lenticellate. Innovations sticky-lacquered. Leaves 
2-3-pinnate; stalks and leaflets articulated; rachis sulcate; leaflets underneath 
minutely but densely dotted, furthermore with gland-fields or scattered glands 
underneath, mostly acuminate, very rarely laxly puberulous. Thyrses terminal, 
very rarely ramiflorous, in one sp. a raceme ; bracts and bracteoles inconspicuous, 
very rarely leafy; thyrses (and calyx) very rarely short-hairy. Flowers white, pink 
with yellow streaks in mouth, more rarely greenish yellow, or golden yellow to 
orange-red. Calyx closed in bud, rarely lobed from the beginning, splitting spatha- 
ceously (not to the base) or mostly irregularly lobed, rarely stunted, often with 
microscopical scale-like glands, moreover mostly with larger crateriform glands; 
after anthesis almost always circumscissile-dehiscing at the base, along an abscis- 
sion line, in one sp. persistent. Corolla either salver- or narrow funnel-shaped or 
more commonly with a mostly short basal tube and often rather suddenly widened 
into an upper tube; lobes mostly rounded unequal, not rarely ciliate. Stamens 
didynamous with a 5th rudiment, but in one Chinese sp. 5 equal stamens, not 
exserted; another-cells V-shaped divergent; filaments inserted at the apex of the 
basal tube, except in two spp. capitate-glandular hairy at the insertion and in the 
basal part, for the rest glabrous; connective produced. Ovary elongate often 
minutely lepidote, glabrous, or tuberculate, never hairy; in both cells with several 
rows of ovules; style filiform, mostly exceeding the anthers; stigma 2-lipped. 
Capsule linear, terete, up to 75 cm; valves smooth, pergamentaceous, rarely ± 
woody, in one sp. tuberculate; septum terete, brittle, corky, but with shallow im- 
pressions of the flat seeds, a narrow line on both sides testimony of attachment to the 
middle of the valves. Seeds very cv>, flat, small, narrow, on both ends with a hyaline 
wing. 

Distr. About 15 spp. in Indo-Malesia, from the DeccantoS. China, Hainan, Formosa, and the southern 
Ryu Kyus, most abundant in tropical SE. Asia; throughout Malesia, but not in the Moluccas proper and 
New Guinea. Fig. 21. 

Ecol. Lowland primary and secondary rain-forests, up to c. 1500 m, not rarely pioneering in disturbed 
forest and on slopes. 

Flower colour and corolla shape vary from pure white to orange-red and from hypocrateriform 
(R. sinica (Hance) Hemsl. and R. frondosa Chun & How) to tubular or campanulate. For the narrow- 
tubular orange-flowered species pollination by birds can be expected (/?. ramiftora) and for the pure 
white, possibly nocturnal ^?. peninsularis hawk-moths may be the pollinating insects. 

Ramifiory is found in Malesia in R. ramiflora; it occurs also occasionally in poor forms of R. pinnata 
after leader-shoots have been damaged. In continental Asia it is also found in R. ignea (Kurz) Steen. 
and in R. hainanensis Mfrk. 

Uses. Minor uses, see under the species. 

Syst. Related to Stereospermum, which differs radically in having only two rows of ovules in each 
cell, and thick trigonous seeds with folded cotyledons fitting in cavities of the septum ; moreover, its leaves 
are always I -pinnate and most species are deciduous and prefer a seasonal climate. 

Two small genera have recently been split off, both from SB. Asia, viz the monotypic Pauhiopia Sin n. 
(Acta Bot. Nccrl. \H, 1969, 425) which has winged rachises, a curved corolla tube, wingless thickish seed, 
a very thin septum, and a truncate calyx open in bud; and Barnettia Saniisuk (Kew Bull. 28, 1973, 172) 
with 2 spp. in Thailand, with I -pinnate leaves in pscudowhorls, a short-ellipsoid, compressed CtfMllfe, 
and tuberculate or immcrscd-glandular calyx and capsule and a cruciform septum; the latter genus 
. relationship with Hrlrrophranrmi 

The closest ally of Rutin mm lurti is obviously the Afro-Asian genus / nmmtloa (< / Smi N. Mlunica 23, 
1976, 133) from which it differs by the terete septum, the usually not ribbed or striate and thinner. 



152 



Flora Malesiana 



[ser. I, vol. 8 2 



narrower fruit valves, mostly 2-4-pinnate leaves, absence of domatia underneath the leaflets, and mostly 
crateriform glands. 

Notes. Specific delimitation, especially in Malesian material, proved difficult, as several species appear 
to be variable and many were only known from the type, notably in the Philippine Islands. Degree of 
pinnation is variable, often in one collection. Also the number and place of the larger glands on under- 
surface of leaflets and on the calyx is liable to variation, as well as the flower colour and the way in which 
the calyx splits. A critical scrutiny showed only few tangible characters and resulted into a severe reduc- 
tion of species in Malesia. 

KEY TO THE SPECIES 

1. Calyx strongly lengthwise 5-6-ridged, l 3 / 4 cm, narrow, one side cleft down ± halfway, 3-toothed at 
apex. Leaves 1 -pinnate, with 5 leaflets, coriaceous, very glossy above, with recurved edge, 7-14 by 
3^4 cm. Corolla 4 cm, with narrow tube, slightly enlarged above; lobes c. 1 cm, obtuse. 

6. R. coriacea 
1. Calyx not lengthwise 5-6-ridged. 
2. Corolla narrowly funnel-shaped, without distinction of basal and upper tube. Flowers in cauli- 

florous racemes, with yellow tube and red limb. Leaves 2-pinnate 1. R. ramiflora 

2. Corolla with a cylindric basal tube, widened rather abruptly into an upper tube. Flowers in terminal 
thyrses which are rarely raceme-like depauperated. 
3. Calyx very short (3-5 mm), with a stunted rim, persistent. Leaves always 1-pinnate 

2. R. glandulosa 
3. Calyx longer, irregularly lobed, after anthesis circumscissile-caducous. 

4. Corolla 2-3 1 / 2 (-4 1 ' 4 ) cm long (incl. lobes). Calyx 5-10C-1 3) mm long 3. R. pinnata 

4. Corolla 5-6 cm long. Calyx 10-25 mm. 
5. Filaments and inside of basal tube glabrous. Corolla tube outside towards apex and on lobes with 
minute sessile glands. Leaves above puberulous on midrib and nerves, beneath with some scattered 

glands. Corolla white or cream-coloured 4. R. peninsularis 

5. Filaments and inside of basal tube near insertions capitate-glandular hairy. Corolla tube outside 
towards apex mostly laxly and very short-hairy. Leaves glabrous above, beneath mostly with at 
least a gland-field at base. Corolla mostly pinkish, sometimes white, with yellow streaks in the 
mouth 5. R. gigantea 



1. Radermachera ramiflora Steen. J. Bot. 72 (1934) 
5;Blumea23(1976) 129. 

Large tree, 24-30 m, stem 30-60 cm ; fluted at 
base; bark grey, fissured. Innovations and racemes 
lacquered, resinous sticky. Leaves (2-)3-pinnate, up 
to 1 m, more or less crowded at the twig-ends ; pin- 
nae 4-6 pairs; leaflets elliptic-lanceolate, acuminate 
at both ends, stalked, 3 1 ' 2 -8 1 / 2 by l 1 '^ 1 2 cm, under- 
neath fine glandular-punctate and with scattered 
small, shallow glands especially near the base, and 
a few scattered flat glandular spots. Flowers thickly 
set in closely placed ramiflorous pendent racemes 
to 20 cm long, erect on curved pedicels 1-2 cm long 
and with 3 bracteoles ± halfway on an articulation. 
Calyx in bud pear-shaped, closed, reddish green, 
tubular, 2-3-lobed, eglandular outside, microsco- 
pically glandular-papillose inside, c. 2V4-2Va cm 
long, 1 cm at mouth, lobes 4-6 mm, 1-2 split ± 
halfway. Corolla with yellow tube and red limb; 
tube slightly curved, narrow salver-shaped, without 
narrowed basal tube, 5-7 cm, the basal 1 1 / 2 cm 
densely pubescent with thick hairs, mouth IV2 - 
2 cm ; lobes rounded, subequal, ± 1 cm 0, ± 
papillose inside. Stamens (and style) reaching the 
mouth, inserted halfway the tube, glabrous but 
their adnate base lax glandular-papillose, thecae — 
divergent, 4-5 mm; no produced connective. Disk 
thick, annular-cup-shaped, faintly crenate. Ovary 
ribbed, pistil c. 4 cm, stigmatic lobes 2 mm, 
very narrow. Capsule straight or twisted, 35-70 cm, 
c. 5 mm 0; septum 2 1 4 -3 mm 0. Seeds 4-5 by 
2V2 mm, the wings 6-7 mm. 

Distr. Malesia: Sabah, Mt Kinabalu (Peni- 
bukan, Dallas, Mesilau, Tenompok, Kota 
Belud, Kp. Kiau I resthouse), not rare. 



Ecol. Rain-forest, also in disturbed forest on 
hill side, 950-1 500 m. Fl. Aug., Jan.-March, fr. 
April, Dec. 

Notes. A characteristic, isolated species: rami- 
florous, flowers in racemes, shape of corolla. 

Leaflets of suckers and saplings dentate. Noote- 
boom & Aban 1603 has only 2-pinnate leaves, 
Clemens 30364 has them 2-3-pinnate. 




Fig. 21. Range of the genus Radermachera Z. & M. 



2. Radermachera glandulosa (Bl.) Miq. Ann. Mus. 
Bot. Lugd.-Bat. 3 (1867) 250; K. & G. J. As. Soc. 
Beng. 74, ii (1905) 380; Koord. Atlas Baum. 2 
(1914) t. 356; Steen. Thesis (1927) 965; Bull. Jard. 
Bot. Btzg III, 10 (1928) 241 ; Corner, Ways. Trees 
(1940) 168, f. 43 ; Chatterjee, Bull. Bot. Soc. Beng. 



1977] 



Bignoniaceae (van Steenis) 



153 



2 (1948) 74; STEEN. Acta Bot. Need. 2 (1953) 307; 
Back. & Bakh./. Fl. Java 2 (1965) 541 ; Santisuk, 
Thai For. Bull. Bot. 8 (1974) 27, f. 15; Steen. 
Blumea 23 (1976) 126. — Spathodea glandulosa 
Bl. Bijdr. (1826) 762. — Bignonia porteriana 
Wall, ex DC. Prod. 9 (1845) 165. — R. stricta 
Z. & M. ex Zoll. Syst. Verz. 3 (1855) 53 ; Ridl. Fl. 
Mai. Pen. 2 (1923) 550. — Stereospermum glandu- 
losum Miq. Suppl. (1860) 240, 565; Clarke, Fl. 
Br. Ind. 4 (1884) 383; K. & V. Bijdr. Booms. 1 
(1894) 74; Kanjilal & Das, Fl. Assam 3 (1939) 
404. — Lagaropyxis glandulosa Miq. Ann. Mus. 
Bot. Lugd.-Bat. 1 (1864) 199. —Fig. 20e-h. 

Small, glabrous, evergreen, crooked tree, up to 
12 m, 40 cm 0; bark slightly pimply and peeling, 
but not flaky or fissured, bitter; young leaves deep 
purple, acrid. Leaves 1-pinnate; leaflets 2-5 pairs, 
large, chartaceous, elliptic (mostly broad-), 
rarely oblong-lanceolate, short-tipped, with a 
conspicuous dark, dense gland-field at the oblique 
base underneath (often bulging on upper surface), 
10-30 by 5-17 cm. Thyrse narrow, 6-50 cm, gradu- 
ally elongating, sometimes the upper part still 
flowering while lower are in fruit. Calyx 3-5 mm, 
cup-shaped, without abscission line, persistent, 
truncate, purple, spotted with 5-7 purple glands in 
a crescent. Corolla narrow, slender, tube narrow, 
slightly curved, halfway rather gradually widening, 
3(-4) cm, pinkish purple outside, white inside, 
with a gland-field at outside of the ciliate lobes, 
basal part of lower tube short capitate-glandular 
hairy inside. Stamens hairy at insertion. Pods 
hanging in bunches, straight, 15-30 cm; valves 
5-7 mm wide; septum 3-4 mm 0. Seeds 10-16 by 
1' 2 -2 mm. 

Distr. Continental SE. Asia (Assam, Burma, 
Thailand, Laos, China: Kwangsi, Kwantung) and 
West Malesia: Malay Peninsula (also Penang), 
Sumatra, Krakatao, West to East Java. 

The two records from Borneo are erroneous: the 
Korthals specimens are mislocalized and Beccari 
811 (mentioned by Merrill, En. Born. 1921, 525) 
is a PS number. Both are from Sumatra. 

Ecol. Primary and secondary forests and thick- 
ets, frequently by streamsides, even rocky 
Saraca-slreams (Corner), both under everwet 
and under seasonal conditions (in Central and 
East Java), from sea-level to 900 m, once reported 
from 5000 ft (Cameroon Highlands). Fl. April, 
July-Nov.,/r. Jan. -Dec. 

The fine seed would indicate easy dispersal, but 
though the species is found in Penang and Krakatao 
Is., it has never been collected in the Riau Is., not 
in the islands west of Sumatra and also not in 
those close to East Java (Madura I., Bali, and 
Kangean) though it is found in Java as far east as 
Bali Straits and R. gigantea extends east as far as 
Alor. The glands at the leaflet basis arc often black 
from sooty moulds indicating actual glandular 
excretion. 

Uses. The timber is small and of ncgligcablc 
value. 

Vcrn. Tuwigadang, M, Minangkahau; ktabako, 
ki hapit, ki lunghit, kipahii, ki sakut, ki uku/>, S; 
ambal, hangkmg, hangkongan, djilihru, godong 

amhoi, harnbal, kawuk rabret, kltjju, lumbal, 

padali, prdali, pudang, J ; sekar poti, Md. 

3. Kackrmachcra pinnata (Hi am o) Sum. J. Bot. 8 



(1870) 147; Merr. Philip. J. Sc. 3 (1908) Bot. 336, 
incl. var. glabra Merr. I.e.; Sp. Blanc. (1918) 350; 
En. Philip. 3 (1923) 446; Steen. Thesis (1927) 973; 
Bull. Jard. Bot. Btzg III, 10 (1928) 248; Blumea 23 
(1976) 129. — Millingtonia pinnata Blanco, Fl. 
Filip. (1837) 501. — Millingtonia quadripinnata 
Blanco, I.e.; ed. 3, 2 (1878) 286, t. 252. — R. 
banaibana Bureau, Adansonia 2 (1861) 194. — 
R. quadripinna Seem. J. Bot. 8 (1870) 147. — Stereo- 
spermum pinnatum F.-Vill. Nov. App. (1880) 151 ; 
Rolfe, J. Linn. Soc. 23 (1884) 314; Vidal, Phan. 
Cuming. Philip. (1885) 132; Rev. PI. Vase. Filip. 
(1886) 203. — Stereospermum quadripinnatum 
F.-Vill. Nov. App. (1880) 151 ; Vidal, Syn. Atlas 
(1883) 35, t. 73. — Stereospermum banaibanai 
Rolfe, J. Linn. Soc. 23 (1884) 314. — Stereo- 
spermum seemannii Rolfe, I.e. — R. acuminata 
Merr. Philip. J. Sc. 3 (1908) Bot. 335; En. Philip. 3 
(1923) 445; Steen. Thesis (1927) 980. — R. mindo- 
rensis Merr. Philip. J. Sc. 3 (1908) Bot. 338; En. 
Philip. 3 (1923) 446; Steen. Thesis (1927) 971. — 
R. fenicis Merr. Philip. J. Sc. 3 (1908) Bot. 335, 
434; En. Philip. 3 (1923) 446; Steen. Thesis (1927) 
979, incl. var. acuminata Steen. ; Jard. Bot. Btzg 
III, 10 (1928) 261. — R. whitfordii Merr. Philip. J. 
Sc. 7 (1912) 352; En. Philip. 3 (1923) 447; Steen. 
Thesis (1927) 963. — R. brachybotrys Merr. 
Philip. J. Sc. 26 (1923) 489; Elmer, Leafl. Philip. 
Bot. 10 (1939) 3809. — R. sorsogonensis Elmer ex 
Steen. Thesis (1927) 973; Elmer, Leafl. Philip. 
Bot. 10(1939) 3809. — R. elegans Steen. Bull. Jard. 
Bot. Btzg III, 10 (1928) 252, f. 8. — R. fenicis (non 
Merr.) Steen. I.e. 261, f. 11. 

See for other synonyms under ssp. acuminata. 

Distr. Malesia: Sumatra (also Simalur, Banka, 
and Siberut Is.), Malaya, Borneo, Philippines, 
Celebes (also Muna I.), and W. Moluccas (Sula 
Is. : Taliabu). 

Notes. It has appeared impossible to separate 
the material into smaller species, as there is 
gradual transition of the many populations, 
especially in the Philippine islands, notably in 
vegatative characters. R. brachybotrys is merely a 
depauperate ridge facies. In one specimen 1- and 
2-pinnate or biternate leaves are not seldom found 
together. For brevity I have omitted from the 
synonymy the many pro parte citations under 
R. amoena, hypostictum and gigantea. Some speci- 
mens from Borneo and Celebes show a tendency in 
leaf-shape towards ssp. acuminata. 

KEY TO THE SUBSPECIES 

1. Leaflets usually chartaceous, elliptic, acuminate 
to caudate; basal gland field mostly distinct, at 
apex eglandular or with a few scattered glands 

.\.\/>. pinnata 

I. Leaflets firmly chartaceous to coriaceous with 
consequently less marked prominent venation 
beneath, usually obovate to elliptic-obovate, at 
apex mostly short- and blunt-tipped or blunt or 
rounded; basal gland field denM, often One also 
at apex ssp. acuminata 

up. pinnata. 

[Yet tO 20m, 15-40 Cn) Z. leaves (1)2-3- 

pinnate, 25 S0( 70) cm; leaflets elliptic-oblong, 

acuminate, mostly at both ends, to caudate, 
(3 )5 I6by(l ! /j )2 5( H)cni. usually chartaceous, 



154 



Flora Malesiana 



[ser. I, vol. 8 2 





—^-^x 




I* 7 




• 


WJ 


J 


91k. 


^Sfl^, 




i3 










r 




.* B ^« 


gs 




v 


- 


< 


j4 



Fig. 22. Radermachera pinnata (Blanco) Seem. ssp. acuminata (Steen.) Steen. Habit, in flower, Aug. 1970 

(photogr. B. C. Stone). 



with a basal gland field, apical gland field absent or 
of scattered glands. Thyrses glabrous, sometimes 
puberulous, (3-6-) 15-60 cm, in odd specimens 
sometimes on the old wood. Calyx l U-\(-\ l l{!) cm, 
lobes unequal, (1— )2— 4, glabrous, usually glandless. 
Corolla pink to pale purplish, with yellow mark- 
ings in the throat, 2-3 1 /2(-4 1 /2) cm, rarely slightly 
lax short-hairy. Stamens glandular-hairy at their 
insertion. Ovary and style glabrous, 12— 17(— 22) 
mm. Capsule (6-)30-50 cm ; valves 4-6(-7) mm 
wide; septum 2-2V 2 rnm 0. Seeds (7— )13— 15 by 
2-3 mm. 

Distr. Malesia: Philippines (the most common 
species), Celebes (also in Muna I.), Moluccas 
(Sula Is. : Taliabu). 

Ecol. Lowland and montane primary and 
secondary forests, on streamsides, up to 600 m. 
Fl. Aug.-May, fr. Febr.-Nov. 

Uses. Minor uses only, for carving and fuel. 

Vern. Philippines: banai-bdnai, Tag., the com- 
mon name, kalapuing, salai, tuing-hulo, ulimbabon, 
yabang-ydbang, Tag., banoi-bdnoi, Bag., ansohan, 
badlan, Bis., labayanan, C.Bis., paling-udk, Bik., 
pagalayan, Bon., bunglai, Buk., atiatip, Ig., 
lanunisi, lasilak, Ibn., barangauan, Ilk., bunlai, 
Mbo,. banaibayan, paitan, pata, Pang., bani-bdni, 
Sbl., kutokong, Sub., hali-hdli, Sulu; Celebes: 
ririh, Muna. 

Note. In a few specimens some inflorescences 
are ramiflorous, obviously due to damage of the 
leader shoot. 

ssp. acuminata (Steen.) Steen. Blumea 23 (1976) 
129. — Spathodea lobbii T. & B. Nat. Tijd. N. I. 



25 (1863) 413. — R. lobbii Miq. Ann. Mus. Bot. 
Lugd.-Bat. 3 (1867) 250; Seem. J. Bot. 8 (1870) 
147; Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 243, 
f. 5, inch ssp. acuminata Steen. I.e. 247, f. 6; 
Corner, Ways. Trees (1940) 168, f. 43 { i lobbiana , )\ 
Santisuk, Thai For. Bull. Bot. 8 (1974) 29. — 
R. amoena [non (Wall.) Seem.] Gamble, J. As. 
Soc. Beng. 74, ii (1905) 381 ; Ridl. Fl. Mai. Pen. 2 
(1920) 551. — R. corymbosa Steen. Bull. Jard. Bot. 
Btzg III, 10 (1928) 249, f. 7. — R. gigantea [non 
(Bl.) Miq.] Burk. Diet. (1935). — Fig. 22. 

Tree, 7-40 m, 60 cmT 0. Leaves biternate or 
2-pinnate; leaflets rather coriaceous, obovate to 
obovate-elliptic, not or short- and blunt-tipped, 
exceptionally acuminate, basal gland field well- 
developed, apical one usually distinct, 8-15 by 
3V2-7V2cm. Thyrse apical, fairly narrow, 15-25 
cm. Calyx 1 cm, rarely to l'^on. Basal tube of 
corolla rather suddenly widened. 

Distr. S. Peninsular Thailand (Krabi, Pattani); 
in Malesia: Sumatra (also in Banka; Mentawai 
Is.: Siberut; Simalur I.), Malaya, Borneo, SW. 
Philippines (Palawan, Culion). 

Ecol. Primary and secondary forests, also in 
open grasslands. In Malaya especially by streams 
(Corner), on granite as well as on limestone; at 
0-400(-800) m. Fl. May, July-March, fr. May- 
No v. 

Taxon. Formerly often accepted as conspecific 
(also by myself, 1927) with R. gigantea cq. amoena, 
but certainly distinct. 

Vern. Sumatra: kaju singamba, sindur langit, 
sundur langit, Batak, kudo kudo pajo, Simalur, 
mentu, tuih, tuwi(k), Banka; kapung suwi, kika- 



1977] 



Bignoniaceae (van Steenis) 



155 



pung, Lampong; bunga pawang, setengah burong, 
tangkani, Malaya; Borneo: binutan, kudjuk langit, 
Dajak-Kapuas; Culion: totancola, Tagb. 

Note. I must admit that I have somewhat 
hesitantly kept this apart as a subspecies from 
true R. pinnata, from which it differs only in shape 
and texture of leaflets and geographical range. 
There are some specimens in Borneo which seem 
transitional. 

In the Philippines (Palawan and Culion) aber- 
rant specimens are found with biternate leaves and 
coriaceous leaflets with recurved margin and rather 
prominent veins underneath. They were collected 
in grasslands which may account for their habit. 
However, in the same islands there are also larger- 
flowered specimens which I have referred to R. 
gigantea with similar habit. More field work in 
these islands is needed to check my tentative 
conclusions. 

4. Radermachera peninsularis Steen. Blumea 23 
(1976) 128, f. la-d. — R. borii {non Fischer) 
Santisuk, Thai For. Bull. Bot. 8 (1974) 30. — 
Fig. 20a-d. 

Tree, 4-15 m. Leaves 3-pinnate, 6O-80cm; 
leaflets lanceolate-oblong, falcate-caudate, 5-8 by 
2-3 cm, on both surfaces microscopically punctate 
but only with a few scattered, very small 'larger' 
glands; midrib and main nerves (c. 4-5 pairs) 
puberulous above (as in R. sinica). Inflorescences 
terminal, similar to those in R. sinica, peduncle 
firm, 15-35 cm long, rachis 4-7 cm; full-grown 
pedicels 2 1 2 -7 cm, halfway with 2 decussate linear 
bracteoles c. I 1 2 -3 cm long. Bracts long, linear, 
exceeding the buds, the lowest up to 5 cm long, 
upper ones 2'/ 2 -3 cm. Calyx campanulate, rather 
wide and thickish, densely microscopically lepi- 
dote, 2-2' 2 by 1 ' 2 cm, rather irregularly lobed for 
1 j- 2 5 of its length, with 5 ± distinct gland fields. 
Corolla white or creamy, c. 6-7 cm long (incl. the 
?entire lobes), with a rather wide ( 3 4 cm) basal 
tube c. 1 ' j cm long concealed in the calyx on top of 
which the filaments are inserted ; upper tube funnel- 
shaped widened. Filaments c. 3 cm, glabrous; 
anthers 4' 2 mm; connective appendage small. 
Ovary glabrous. Capsule terete, tortuous, 60-70 cm 
by 3-5 mm. 

Distr. Peninsular Thailand (between Phangnga 
and Krabi);in Malesia: Malay Peninsula (Cameron 
Highlands), 2 collections. 

Ecol. In Peninsular Thailand at 8 25' N, 99 15' 
E, in evergreen forest along a stream, on limestone, 
at 50 m altitude, in the Cameron Highlands at 
1200 m in mixed rain-forest. 

Notes. In habit deceptively like R. sinica, but 
at once different by the shorter (6-7 cm), not 
salver-shaped corolla and the much wider smooth 
calyx. The corolla in R. sinica measures c. H 12 cm 
including the lobes, with a narrow, very gradually 
widening tube. R. sinica also has the minute lax 
pubescent nerves and veins above and long narrow 
bracts. 

SANTW k I.e. referred the specimens to R. borii, 
which I refer to R. unua, a species ranging more 
northerly in SE. Asia, viz from N. Assam, N. 
Burma, and Tonkin to S China, Hainan, 1 ormoaa, 
and the southern Ryu Kyu I 

5. Radermachera gistantea (Mi.) Mio. Ann. Mus. 



Bot. Lugd.-Bat. 3 (1867) 250; Seem. J. Bot. 8 
(1870) 146; Koord. Atlas Baum. 2 (1914) t. 356 
A-K; Heyne, Nutt. PI. (1927) 1371 ; Steen. Thesis 
(1927) 983, p.p., excl. syn. lobbii; Bull. Jard. Bot. 
Btzg III, 10 (1928) 253, f. 9; Blumea 23 (1976) 126. 

— Bignonia gigantea Norona, Verh. Bat. Gen. 
5 (1790) 70, nomen. — Spathodea gigantea Bi_. 
Bijdr. (1826) 761 ; Miq. Fl. Ind. Bat. 2 (1858) 751. 

— Bignonia amoena Wall. [Cat. (1832) n. 6512, 
nomen) PI. As. Rar. 2 (1831) 78, t. 183; Loudon, 
Hort. Brit. (1830) 483, err. amara; G. Don, Gen. 
Hist. 4 (1838) 222. — Bignonia oxyphylla DC. 
Prod. 9 (1845) 169. — Stereospermum hypostictum 
Miq. Sum. (1861) 565; Clarke, Fl. Br. Ind. 4 
(1884) 384, excl. syn. S. lobbii; K. & V. Bijdr. 
Booms. 1 (1894) 72, excl. syn. lobbii; Kanjilal & 
Das, Fl. Assam 3 (1939) 405. — Lagaropyxis 
gigantea Miq. Ann. Mus. Bot. Lugd.-Bat. 1 (1864) 
198, incl. f. sumatrana et f. borneensis MlQ. — 
R. amoena Seem. J. Bot. 8 (1870) 146; non Gamble, 
J. As. Soc. Beng. 74, ii (1905) 381, quae est R. lobbii. 

— R. elmeri Merr. Bull. Gov. Lab. Philip. 29 
(1905) 48; Philip. J. Sc. 3 (1908) Bot. 334; En. 
Philip. 3 (1923) 445; Steen. Thesis (1927) 994. — 
R. biternata Merr. Philip. J. Sc. 1 (1906) Suppl. 
238; ibid. 3 (1908) Bot. 333; En. Philip. 3 (1923) 
445; Steen. Thesis (1927) 970. — R. palawanensis 
Merr. Philip. J. Sc. 3 (1908) Bot. 336; Steen. 
Thesis (1927) 977. — R. elliptica Merr. Philip. J. 
Sc. 3 (1908) Bot. 334; En. Philip. 3 (1923) 445; 
Steen. Thesis (1927) 964. — R. sibuyanensis Elmer, 
Leafl. Philip. Bot. 4 (1912) 1485; Merr. En. Philip. 
3 (1923) 447; Steen. Thesis (1927) 992. — R. elmeri 
var. fragrans Elmer, Leafl. Philip. Bot. 7 (1915) 
2561; Merr. En. Philip. 3 (1923) 446. — R. 
fragrans Steen. Thesis (1927) 996. — R. punctata 
Elmer ex Steen. Thesis (1927) 982; Elmer, Leafl. 
10 (1939) 3709. — R. borneensis Steen. Bull. Jard. 
Bot. Btzg III, 10 (1928) 258, f. 10. 

Shrub or tree, (6-)20-40 m, up to 80 cm ; bark 
and young leaves bitter. Leaves (l-)2(-3)-pinnate, 
12-35(-80) cm, leaflets usually elliptic to oblong, 
rarely somewhat obovate or lanceolate, shorter or 
longer acuminate, 4— 1 2(— 1 5 ) by 2-6(-9) cm, at base 
underneath mostly with a gland-field, at apex with 
some scattered glands. Thyrses 8-40 cm, rather 
open, terminal, cns -flowered, glabrous. Flowers not 
rarely fragrant. Calyx <1— >1 l /^— 2 1 ; 2 cm long, 
mostly 2-lobed, sometimes 1 - or more-lobed ; glands 
few or distinct. Corolla 5-6 cm long (incl. lobes), 
pink or white, usually with yellow streaks in the 
mouth, above the basal tube rather suddenly 
widened, campanulate upper tube towards apex 
almost always short-capitate glandular hairy, and 
lobes ciliate. Filaments densely capitate-glandular 
hairy at insertion. Style 2' 2 3 cm. Capsule 15-60 
cm long, 5-8 mm <Z ; septum 4 mm 0. Seeds 8-13 
by 2-4 mm. 

Distr. SE. Asia (Assam: Khasi & Jaintia Hills; 
Burma: Manipur, Tavoy, Wallich, Clarki, I.e., 
not seen), and Malesia Sumatra (incl. Billiton and 
Hanka), West to East Java (common in Central 
and 1 ast Java), I csser Suiul.i Is. (Bali, Sumhawa. 
\ lores, Alor), Baucan I , SE Borneo, and Philip- 
pines. 

Ecol. Primary and secondary forests, also in 
areas subject to a dry monsoon, and in East Java 
in teak forest, very common on Mt Telemojo and 
near Pringombo (Banjumas) (Koorders, I.e.), 



156 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 23. Fernandoa macroloba (Miq.) Steen. a. Habit of branchlet with young fruit, x 1 / 2 , b. inflorescence 

in bud, x 1 j 2 , c. calyx, nat. size, d. corolla, x 3 / 4 , e. CS of young capsule, nat. size, f. mature capsule, 

x 1 j 2 , g. seed, nat. size, h. detail of underside of leaflet, showing small domatia and flat gland, x 4, (a-e, 

h C.H.B. XI-C-106a,/-# Diepenhorst 2353 HB, type). 



1977] 



Bignoniaceae (van Steenis) 



157 



from the lowland up to c. 1600m. Fl. Jan-Dec, 
fr. Jan.-Dec. Not rarely a pioneer in secondary 
forest, cuttings sometimes used for strengthening 
terracing on unstable slopes. Early flowering and 
fruiting as a shrub. 

Uses. Sometimes cultivated; also suitable for 
re-afforestation. Timber is said to be strong and 
durable, locally used for bridges and houses, but 
not resistent against termites, and therefore of less 
value for outdoor constructions ; used for making 
matches and matchboxes (Heyne, I.e.). 

Vern. Sumatra: tui (batu), M, Billiton, 
Minangk., djamatan, djamaton, kaju angin, kaju 
diling, radja matan, simaisaludang, sunder langit, 
M, Karo-Batak, tangke, M, Atjeh, kulit berilang, 
M, Kroei, (ke)kapung tut, talas, M, Lampong; 
West Java : kipadali, S ; East Java : bedali (common 
name), kedah, (ke)dali, kokok-kejok, J, kaju raras, 
karpoteh, potian, putian, sekar petak, sekar pote, 
Md; pedanten, Bali; atodjang, Alor; Borneo: 
binutan, Dajak, Martapura, bunglai batu, b. 
gunung; Philippines: Palawan: agtap, tantangan, 
Tagb., sayo, Ig., barangau-a-nalabdga, pamaya- 
bayen, Ilk. 

Notes. I have come to the conclusion that the 
large-flowered species of the Philippines, R. 
borneensis and R. gigantea, should be referred to 
one variable taxon. Though there may be local 
races they can not be properly distinguished, not 
even on subspecific level, as their 'characters' fade 
away; R. elliptic a with 1 -pinnate leaves goes via 
biternate leaves to 2-pinnate leaves. R. sibuyanensis 
has an almost glabrous corolla; R. fragrans has 
fragrant flowers, but fragrancy is mentioned 
frequently on labels, but also sometimes flowers are 
said to be scentless. Flower colour is said to be 
white or pink in the Philippines, to which mostly is 
added the occurrence of yellow streaks in the 
mouth. White flowers are only reported from 
Borneo and the Philippines. 

Especially a few Philippine specimens from Pala- 
wan and Culion are aberrant; earlier collections 
were described by Merrill as R. palawanensis and 
R. biternata. They were collected in grasslands and 
differ by thicker, often biternate leaves, leaflets 
with recurved margin and often distinctly arching 
prominent veins on the undersurface. One number 



(PNH 12319) has singularly slender tubular flowers 

5 cm long. Other specimens with the same vegeta- 
tive difference from normal R. gigantea have, how- 
ever, smaller flowers (3-3 V2 cm) and these I have 
identified as R. pinnata ssp. acuminata. I must admit 
that closer field work is needed to check my tenta- 
tive conclusions. 

R. amoena (Wall.) Seem, was described from a 
flowering shrub in Hort. Calc. which Wallich 
found worthy of an ornamental. Wallich was 
somewhat doubtful about its provenance, but 
assumed that it was introduced by Mr C. Telfair 
from Mauritius, where it must then also have been 
cultivated. Though it is not known from Malaya 
and Thailand, it does occur in Assam (Khasia & 
Jaintia Hills, cf. Kanjilal & Das) and N. Burma 
(Manipur: Meebold 5169). These specimens I can- 
not distinguish from R. gigantea. What Kanjilal 

6 Das mean in their key by 'rusty-coloured cap- 
sules' is not clear to me ; they mention the flowers to 
be white. 

Insufficiently known 

6. Radermachera coriacea Merr. Philip. J. Sc. 3 
(1908) Bot. 333; En. Philip. 3 (1923) 445; Steen. 
Thesis (1927) 961 ; Blumea 23 (1976) 131. 

Leaves 1-pinnate, 20-30 cm; leaflets 5, coria- 
ceous, oblong to elliptic-oblong, base acute, apex 
blunt or obscurely blunt-tipped, margin recurved, 
7-14 by 3-4 cm, very glossy above; nerves 13 
pairs. Panicles 15 cm. Calyx l 3 / 4 cm, narrow, 
strongly lengthwise 5-6-ridged, cleft on one side 
halfway, 3-toothed at apex. Corolla 4 cm, the tube 
rather narrow, slightly enlarged above, the lobes 
obtuse c. 1 cm. Capsule 16 cm, valves 5-7 mm wide 
{ex descr.). 

Distr. Malesia: Philippines: Central E. Luzon 
(Tayabas Prov. : Baler), only known from the type 
(Merrill 1099); vern. name bibit parang. 

Note. This is the only specific name in the genus 
of which I could not trace the type, which may be 
lost. A lengthwise ridged calyx 1 have not observed 
in any species of the genus; the 1-pinnate leaves 
and narrow leaflets seem rather characteristic. It 
might possibly be an extreme form of R. pinnata 
ssp. acuminata. 



10. FERNANDOA 



Welw. ex Seem. J. Bot. 3 (1865) 330, t. 37-38, sphalma Ferdirumdia; ibid. 4 (1866) 
123; ibid. 8 (1870) 280, sphalma Ferdinandoa\ ibid. 9 (1871) 81 ; Milne-Redhead, 
Keu Bull. 3 (1948) 171 ; Heine, Adansonia 4 (1964) 467-^70; Gentry, Ann. Mo. 
Bot. Card. 62 (1975) 480; Shin. Blumea 23 (1976) 133. — Kigelianthe Baill. Hist. 
PI. 10 ( 1891 ) 50. Haplophragma P.Dop, Bull. Soc. Bot. Fr. 72 (1925) 889; Steen. 
Thesis (1927) 998; Bull. Jard. Bot. Btzg III, 10 (1928) 262. -- Spathodeopsis 
P.DOP, C R. Ac. Sc. Paris 189 (1929) 1097. - Hexaneurocarpon P.Dop, I.e. 
Tisseranthodendron Su i ans. Bull. Soc. Bot. Fr. 98 (1951) 270-272, pi. — Fig. 23, 
25. 

Trees. Leaves I -pinnate; leaflets 2-5 pairs, diminishing in si/e downwards, ter- 
minal one largest, beneath glabrous or tomentose with stellate hairs, with few 



158 Flora Malesiana [ser. I, vol. 8 2 

scattered, larger, flat, spot-like glands and small hairy domatia in the nerve-axils. 
Thyrses axillary or terminal, densely or laxly tomentose, or at least almost always 
with (sometimes) small and appressed stellate hairs. Calyx tubular or campanulate, 
unequally 2-5-lobed, with few to many warty or prominent glands in the upper 
half. Corolla with a fairly large basal tube, rather suddenly campanulate-funnel- 
shaped widened to the mouth; lobes undulate to crenate. Stamens 4, didynamous; 
5th rudimentary; anthers divaricate. Disk annular. Ovary elongate, with 2 mar- 
ginal placentas in each cell; ovules oo. Capsule long, linear, terete, twisted or 
straight, pendent, smooth or ribbed; septum flat, smooth, not rarely glossy, 
thickish. Seeds numerous, rather rectangular, the wings rather narrow; insertion 
punctiform. 

Distr. About 4 spp. in tropical West Africa (Angola to Gabon) and East Africa (Tanganyika), 3 in 
Madagascar, and 6 in Indo-Malesia (India to Indo-China); in Malesia 2 spp., in the Malay Peninsula and 
N. Sumatra. Fig. 24. 

Ecol. Tropical lowland forest. 

Taxon. In Blumea I.e. I have given the reasons for uniting all these genera, by which the range of 
Fernandoa is considerably extended and becomes similar to that of Stereospermum, Dolichandrone and 
Markhamia. It differs from Markhamia and Dolichandrone in not having a false septum and a spathaceous 
calyx; it differs from the continental Asian genus Bamettia in not having a cruciate septum, shortly 
ellipsoid-oblong capsules and pseudo-whorled leaves. Its closest ally is Radermachera which has thinner, 
linear-terete capsules which are not striate or ribbed and with thinner valves (except in R. sinica), and a 
terete septum; moreover, the leaves in Radermachera are mostly 2-4-pinnate and the leaflets have no 
domatia and another type of glands and never stellate hairs. 

It is noteworthy that Fernandoa adenophylla shows in the aspect of flower size and shape, thick indu- 
ment and especially in the leaves (leaflet texture and size diminishing downwards, often auricles) a 
striking similarity in habit with Markhamia {e.g. M. cauda-felina), a genus which may have also ridged 
pods in Africa : but Markhamia has a 4-angled septum and a spathaceous calyx ! One gets the impression 
that there are signs of reticulate affinity among the Afro-Asian genera, and possibly parallel evolution, 
but it must be observed that several genera are obviously not unnatural in having distinct pollen types. 

From its wide distribution Fernandoa might be supposed to be an ancient genus, from which Rader- 
machera, Barnettia Santisuk (cf. Kew Bull. 28, 1973, 172) and Stereospermum are derived specialisations. 

Uses. All Indo-Malesian species seem to be good timber trees and of the Indo-Chinese species it is said 
that the timber is good for all purposes and not attacked by termites. They may already flower and fruit 
at an early age. 

KEY TO THE SPECIES 

1. Leaves hairy beneath. Inflorescence terminal, rusty tomentose, with stellate and branched hairs. 
Calyx thick, campanulate, 2V2-3V2 cm, woolly-tomentose outside. Corolla thick, 6-7 cm, whitish to 
yellowish. Capsule twisted, rusty-tomentose, 10-ribbed 1. F. adenophylla 

1. Leaves glabrous except a few short hairs of the domatia. Inflorescences lateral, with appressed, fine, 
partly stellate hairs. Calyx tubular, thin, c. I-IV4CIT1 by 3 mm, fine appressed stellate-hairy. Corolla 
thin, pink, short-hairy outside, c. 4-5 cm. Capsule straight or ± twisted, glabrous, smooth. 

2. F. macroloba 

1. Fernandoa adenophylla (G. Don) Steen. Blumea with irregular crown, 10-35 cm ; innovations, 

23 (1976) 135. — Bignonia adenophylla [Wall, thyrse and calyx with dark rusty, multicellular 

Cat. 6502] ex G.Don, Gen. Syst. 4 (1838) 221. — compactly branched hairs. Leaves 20-50 cm long; 

Spathodea adenophylla DC. Prod. 9 (1845) 206; leaflets 2-3(-4) pairs, subsessile, the lowest often 

Wight, 111. 1 (1839) t. 160. — Heterophragma close to the base of the petiole and smallish and 

adenophyllum Seem, ex B. & H. Gen. PI. 2 (1875) orbicular auricle-like, the terminal leaflet largest, 

1047; Kurz, Fl. Burma 2 (1877) 236; Clarke, Fl. mostly obovate to oblong to elliptic, obtuse to 

Br. Ind. 4 (1884) 381; Prain, J. As. Soc. Beng. acuminate, underneath with scattered crateriform 

60, ii (1891) 322; Koord. Ann. Jard. Bot. Btzg 14 plate glands and glandular spots and with yellow- 

(1897) 417; Gamble, Man. (1902) 514; Ridl. Fl. ish-pubescent, stellate hairs (sometimes on short, 

Mai. Pen. 2 (1923) 551. — Haplophragma adeno- multicellular stalks), 7-24 by 4-19 cm; petiole 

phyllum (DC.) P. Dop, Bull. Soc. Bot. Fr. 72 0-8 cm, sulcate as is the rachis. Thyrse terminal, 

(1925) 890; Steen. Thesis (1927) 1006, f. 13a, stout, erect, lax-flowered, c. 20 cm. Calyx cam- 

14e-f; Bull. Jard. Bot. Btzg III, 10 (1928) 265; panulate, c. 2 l l 2 -3 l l 2 by F^cm, inside sordid 

Gentry, Ann. Mo. Bot. Gard. 60 (1973) 856, f. 14. white, thick, persistent; lobes V2-I cm, subequal, 

Tree, 4-20 m, not rarely poorly developed and entire. Corolla yellow-brown, white, yellowish 



1977] 



Bignoniaceae (van Steenis) 



159 



green (brown in sicco), outside woolly tomentose, 
inside glabrous, basal tube c. l\\-2cm, upper 
tube c. 3^4 cm; the mouth c. 5 cm 0; lobes ± 
entire, l-l 1 2 by 1 ' 2 -2' 2 cm . Anthers ± included, 
cells ± free. Ovary elongate, brown stellate hairy, 
1 cm, style 4 cm, ± exceeding longer stamens. 
Capsule subterete, pendulous, twisted, rusty by 
stellate hairs, 30-60 by l l /2-2 l /2cm 0, the valves 
with 5 strong prominent ribs; septum flat, shining, 
corky, over 1 cm wide. Seeds variable in size in one 
capsule, c. 2 1 2 -3 3 4 by 3 ,' 4 -l V 4 cm incl. the smallish 
wings. 

Distr. Continental SE. Asia (Assam to Tenas- 
serim and Chittagong, Burma, Thailand, Indo- 
China; also in the Andaman and Coco Is. : Prain, 
1891); in Malesia: only in the extreme northern 
part of Malaya (Langkawi, Perlis: Chupeng; 
Kedah: Alor Star; in 1882 also found on Bt. 
Timah in Singapore I.). 

Ecol. Mixed deciduous or evergreen monsoon 
forest, also in bamboo forest often on limestone 
and calcareous soils, but in Burma preferring 
pervious siliceous soils (Kurz) ; 0-750 m. Fl. 
May-Sept.,/r. March-Sept. 

Kostermans (n. 1436) noted that in Thailand 
the 'white' flowers open at night and drop the next 
morning. Kurz I.e. stated it leaf-shedding in 
Burma and flowering at the close of the cold 
season. Van der Pul (Act. Bot. Need. 5, 1956, 
1 38) stated that the nocturnal flowers are visited by 
bats. 



the lowest pair often (2-4 cm) small and roundish, 
14-18 by 4-7 cm; underneath some scattered dark 
glandular spots and minute hairy domatia. 
Thyrses rather short (2-8 cm), lateral, narrow and 
dense, densely appressed pubescent; peduncle 
short; pedicels c. \' 2 cm. Calyx tubular, less than 
halfway with 2-3 incised, short, acute lobes, 
1-1 '/4 cm by 3 mm, appressed-pubescent by 
appressed partly stellate hairs and a few warty 
elevated glands in upper half, faintly 5-ribbed. 
Corolla white or pale pinkish, of thin texture, out- 
side minutely puberulous, 4-5 cm (incl. lobes); 
basal tube c. l 3 / 4 cm long, upper one wide- funnel- 
shaped, c. lV^cm long; lobes unequal obovate- 
roundish, with crenulate margin, 1 1 / 2 — l 3 /* by 
l 3 / 4 -2V 4 cm. Stamens 4, didynamous, inserted 
halfway the basal tube. Disk annular, ± puberu- 
lous. Ovary lanceolate, densely yellow-tomentose, 
4-angular, faintly 6-ribbed; ovules in 6 rows in 
each cell ; style delicate, 1 \' 2 -2 cm, at base ap- 
pressed-hairy. Capsule linear, terete, straight or 
± twisted, glabrous, 40-65 by 1-1 V2 cm 0; 
valves thin-coriaceous, lenticellate ; septum flat, 
3 ' 4 -l cm wide, shining, of corky texture. Seeds 
2-2 x li cm (incl. the very short hyaline wing) by 
3 4 cm. 

Distr. Malesia: northern half of Sumatra: 
from the Res. Minangkabau and Pariaman north- 
ward to Atjeh and even on P. Breueh. 




Fig. 24. Range of the genus Fernandoa Seem. 
Figures above the hyphen indicate endemic species, 
those below the hyphen non-endemic species for 
each area or subarea. In Africa and SE. Asia the 
delimitation is generalized. 



2. Fernandoa macroloba (Miq.) Steen. Blumea 23 
(1976) 136. — Spathodea sp. Teysm. Nat. Tijd. 
V I 14 (1857) 345. — Spathodea macroloba Miq. 
Sum. (1861) 565; Heyne, Nutt. PI. ed. I, 4 (1917) 
167. — Heterophragma macrolobum Ba< k ,\ 
H E, Nutt. PI. cd. 2 (1927) 1371. — Haplo- 
phragma macrolobum (Mia) Sins I hesis (1927) 
1002, f. 13b, I4a-d; Bull. Jard. Bot. Bt/g III, 10 
(1928) 263, f. 12. H«. 23, 25. 

Tall, deciduous tree, 1 5 40 m (bole 15 20 m), 
. no buttresses; larger twigs terete. 
Leaves 25 40cm; leaflets 3 5 pairs, a tew mm 
stalked to sessile, obovatc- (<> elliptic-oblong, 
glabrous, beneath with a few large spot-glands, 
entire, abruptly acute-acuminate, cuncatc at base, 




Fig. 25. Fernandoa macroloba (MlQ.) Sin H. Twig 

with flowers, nat. si/c (photogr. Huysmans, Nov. 

1956). 



160 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 26. Lamiodendron magnificum Steen. a. End of twig, x Vs. b. bud, nat. size, c. ramifiorous raceme, 
d. terminal raceme, both x Vs. e. LS section of ovary, x 3, f. CS of ditto, x 5 (Brass 25543). 



12. PAJANELIA 

A.P.DC. Bibl. Univ. Geneve II, 17 (1838) 130, repr. p. 14; A.DC. Prod. 9 (1845) 
227; Bureau, Mon. (1864) 35, 50, t. 20; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b 
(1895)244. — Fig. 27, 29. 



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Bignoniaceae (van Steenis) 



161 



Ecol. Rain-forest, up to 350 m, according to 
field labels in most localities rather common; see 
e.g. Teysmann, I.e. Fl. Oct.-Dec, fir. Dec-May. 

Uses. An estimated timber tree, used for boats 
etc. (Heyne, I.e.). 

Vern. Tuwe, Atjeh, tuhi, Karo-Batak, radja, 
Batak, sungke (tjirit)), M, Padang Uplands, 
sungkai rimbo, Minangkabau. The name sungkai 
is also used in Palembang for another timber tree 
with opposite pinnate leaves: Peronema canescens 
( Verben.). 



Note. Den Berger & Bianchi (Tectona 24, 
1931, 894-903) noted that the vessels contain a 
sulphur-yellow substance colouring red with alco- 
hol : lapachol ; a rare feature. 

This tree is cultivated in Hort. Bog. sub n. 
XI-C-106A and XI-H-27; unfortunately dupli- 
cates of several other numbers (XI-I-10, 20 & 50) 
have been distributed under this name also; they, 
however, all belong to Dolichandrone serrulata 
(Wall, ex DC.) Seem. 



11. LAMIODENDRON 

Steen. Nova Guinea n.s. 8 (2) (1957) 381, f. 1. — Fig. 26. 

Evergreen tree. Twigs thick, hard, lenticellate, nodes with pitted glands, inter- 
nodes flattened. Leaves 1 -pinnate; leaflets without domatia. Racemes or panicles 
terminal or from the twigs, hairy and with numerous capitate-glandular hairs. 
Thyrses terminal, sometimes reduced to short-racemiform. Flowers large, not 
articulated with the pedicels. Pedicels bracteolate, often with abortive flowers in 
the axil. Calyx halfway split, valvately 5-lobed already in bud, thin, veined. Corolla 
campanulate, hardly zygomorphic, without basal tube, glabrous, glandular-punc- 
tate; lobes 5, imbricate in bud, rounded, entire. Stamens 4 and a staminode, 
inserted in the hairy throat. Ovary 4-sulcate, each cell with 2 placentas on the sep- 
tum, ovules very many, in many rows; stigma with 2 oblong-spathulate lips. 
Capsule (immature, 4 l / 2 cm long) linear, shortly beaked, oval in CS with a flat 
septum on which developed winged ovules; pericarp with 2 fine grooves in the 
middle of the valves and obviously the capsule loculicid. 

Distr. Monotypic. Malesia: E. New Guinea: Milne Bay Distr., d'Entrecasteaux Is. (Normanby I.) and 
Louisiades (Rossel I.). 

Ecol. Coastal lowland forest, rain-forest and sago swamps. 

Taxon. Possibly related to Fernandoa, but lacking domatia and with small crateriform glands on the 
leaflets beneath; the fruit structure may yield further criteria. 



1. Lamiodendron magnificum Steen. Nova Guinea 
n.s. 8 (2) (1957) 381, f. 1 ; Blumea 18 (1970) 563. — 
Fig. 26. 

Tree, 12-20m; flush purplish green. Leaves 
4-5-jugate, glabrous, 30-50 cm; petiole (2'/ 2 -)6- 
10 cm; petiolules 1-2 cm; leaflets firmly charta- 
ceous, ovate-oblong, acuminate, oblique with 
unequal base, 7 1 9 by 3 ' 2 9 cm; nerves 7-1 1 pairs; 
beneath with scattered, small, pitted crateriform 
glands. Thyrsvs glandular-hairy, 2-3 times 
cymosely branched, up to 18 cm, sometimes 
seemingly from old wood reduced to racemiform 
and rachis I 3 cm. Bracts ovate-acute, 3-4 mm. 
Pedicels I' 2 2' .cm, with 1 2 pairs of bracteoles 
3 5 mm long. Calyx 2 3 cm, with a few glands, 
rather densely short capitate-glandular hairy; 
lobes widc-ovatc, mucmnalc, I I ' 2 by ' < I cm. 
Corolla showy, brilliant orange to apricot, dark 
purplc-rcd veined, with many small glands and the 
lobes (and tube) laxly capitate-glandular hairy, 
H 10 by 4' , 6 <_m; lobes l f a 2 cm long, 3 V , un 
wide. Stamens 5 7 cm long, inserted near the base 



of the corolla, at the base densely hairy; staminode 
2 cm; anthers with free, divaricate cells, not versa- 
tile, reflexed, 6 mm long. Ovary quadrangular- 
fusiform, 1 cm long, with large impressed glands, 
moreover covered by dense, microscopical 1 -celled 
hairs and short-stalked peltate to capitate-glandu- 
lar hairs; style 4-6 cm; stigmatic lobes oblong- 
acuminate, 4 mm. 

Distr. Malesia: E. New Guinea (Milne Bay 
Distr., Raba Raba Subdistr., Biniguni, May I track, 
9°38' S, 149 18' E; Northern Distr., Popondetta 
Subdistr., road to Gona-Arunda; near Wanigela; 
near Alotau), Normanby and Rossel Is. 

Ecol. In Normanby I. forming a community on 
a gravel bank behind the beach fronting swamp 
forest, near Wanigela in sago swamps, in Rossel I. 
at 10 m in a rain-forest on a stream bank, near 
Biniguni in poor lowland rain-forest at 90 m, in 
Popondetta in disturbed forest, near Alotau 
( i' i i i wi i i saw ■ single tree. 

Note. Worthy of introduction Into cultivation 
for the large showy flowers. 



162 



Flora Malesiana 



[ser. I, vol. 8 2 



and petiole sharp-keeled above^r^coa, ^ conaC eous, w 




Fi , „. — ^^^z^z^i^^^^' 

d. capsule, e. ditto in CS, both x / 2 , i. ^ rr lg547) 



1977] 



Bignoniaceae (van Steenis) 



163 



± constricted above basal tube, zygomorphous ; lobes 5, spreading to recurved, 
nearly equal, 2 lobes connate halfway up, imbricate in bud. Stamens 4, inserted at 
apex of lower tube, didynamous, subexserted, 5th rudimentary; filaments thick, 
anther-cells divergent. Disk large, annular, fleshy. Ovary oblong-cylindric; ovules 
in each cell cv>, on 2 placentas, cv>-seriate. Capsule flat, obovate-lanceolate, tipped, 
valves broadly winged along the margin. Seeds in several rows on the edges of the 
septum, compressed, hyaline-winged. 

Distr. Monotypic, from Malabar eastwards through SE. Asia (E. Bengal: Khasia, Sylhet; Burma: 
Pegu, Travancore, Chittagong, Tenasserim; Thailand, Andaman Is.) to West Malesia: N. Sumatra (E. 
Atjeh), Malaya (Perak, N. Kedah, ?Singapore, ?Penang), and the Natuna Is. (Sedanau, Bunguran, 
Duperre) in the S. China Sea, NW. of Sarawak. Fig. 28. 

F.-Villar (Nov. App. 1880, 150) recorded it erroneously from the Philippines. 

Ecol. Lowland evergreen mixed forests, in the Andaman Is. also in deciduous forest. 

I assume bats visit the probably nocturnal flowers, but there are no direct records. 



1. Pajanelia longifolia (Willd.) K.Sch. in E. & P. 
Nat. Pfl. Fam. 4, 3b (1895) 244; Steen. Bull. Jard. 
Bot. Btzg III, 10 (1928) 267; ibid. 12 (1932) 164, 
f. 2 (map). — Pajaneli Rheede, Hort. Mai. 1 (1678) 
79, t. 44, sphalma in textu t. "45". — Bignonia 
indica var. B Linne, Sp. PI. (1753) 625. — Bignonia 
longifolia Willd. Sp. PI. 3 (1800) 306. — Bignonia 
pajanelia Buch. Ham. Trans. Linn. Soc. 13 (1821) 
516, nom illeg., quoting Willd. — Bignonia multi- 
juga Wall. [Cat. 6503] PI. As. Rar. 1 (1830) 81, 
t. 95-96; G. Don, Gen. Syst. 4 (1838) 221.— 
P. multijuga A.P.DC. Bibl. Univ. Geneve II, 17 
(1838) 130; A.DC. Prod. 9 (1845) 227, excl. syn 
Lour, et Pers.; Miq. Fl. Ind. Bat. 2 (1858) 758 
Bureau, Mon. (1864) 50, t. 20; Kurz, Prel. Rep 
For. Veget. Pegu (1875) App. A: xciii, App. B: 69 
Fl. Burma 2 (1877) 237 CPayanelia'); Gamble 
J. As. Soc. Beng. 74, ii (1905) 382; Ridl. Fl. Mai 
Pen. 2 (1923) 549. — P. rheedii Wight, Ic. 4 (1848) 
t. 1343-4; 111. 2 (1850) t. 161bis; Bedd. Fl. Sylv. 3 
(1872) clxix, t. 21-5; Clarke, Fl. Br. Ind. 4 (1884) 
384; Brandis, Ind. Trees (1906) 494; Parkinson, 
For. Fl. Andam. (1923) 215. — P. bijuga, lapsus 
mini, in syn. Bull. Jard. Bot. Btzg III, 10 (1928) 
267. — Fig. 27, 29. 

Evergreen, small to large tree up to 30 m, 1 m 0, 
changing leaves before flowering. Innovations 
resinous. Twigs terete, thick, lenticellate. Leaves 
1-pinnate, 8-12-jugate, 40-120 cm; petiole thick, 
to 15 cm, rachis sharply keeled above, rounded 
beneath; leaflets entire, oblique, ovate-oblong, 
tipped, chartaceous, 10-20 by 5-8 cm, 3-5 mm 
stalked; nerves anterior side 9-12, posterior 
6-9 pairs; beneath with scattered or heaped 
crateriform glands along the midrib. Thyrses up to 
1 m, coarse, rachis hollow, 1 cm ; pedicels 
2' | cm; bracteoles minute. Flowers with a soapy 
smell, pale yellowish, inside dull-purple shaded. 
Calyx 3 5 cm, densely covered with microscopical 
glands and besides with scattered, rimmed, dish- 
shaped, large glands; splitting into 5 irregular, 
crispatcly-crcnulatc, acute lobes, tube at base with 
keel-like folds, persistent. Corolla 5 7'/ 2 cm, thick, 
constricted above base, pubescent except at 
base; lobes imbricate in bud, in anthesis reflexed, 
broad-obovatc, crcnulatc and crispatc; anthers 
brown to nigrescent, cells divergent. Style long, 
with a 2-lobed, clavatc stigma. Capsule stiped, 



30-45 by 5-9 cm (incl. the 2-3 cm wide, often 
splitting wings), with a dorsal ridge, and a corky 
margin on which the wings; septum quadrangular 
in CS, contracted in the middle, corky. Seeds in 
oo rows on each margin of the septum, curved, 
274-3 by 3 / 4 -lV4cm (incl. the hyaline wings). 

Distr. As the genus. Erroneously recorded for 
the Philippines by F.-Villar, Nov. App. (1880) 
150. Rarely planted (Atjeh). Fig. 28. 




Fig. 28. Range of the genus Pajanelia A.P.DC. 



Ecol. Lowland primary and secondary rain- 
forest, in the Natuna Is. common, with plenty of 
seedlings, spared in coconut stands, elsewhere 
scattered, riverbanks, etc., mostly coastal, from 
sea-level to 100 m ; in the Ghats recorded to 700 m 
altitude. Fl. Jan. -April, Aug., fr. March-April, 
Aug. 

Indian authors cite it as being evergreen, but in 
the Natuna Is. I found it shortly deciduous. Kurz 
found it in Burma an evergreen tree. He recorded it 
from the Andaman Is. as 'very abundant in the 
leafless jungles', that is: monsoon forest (Rep. Veg. 
Andam. Is. 1870, 12, 43, 71). Parkinson gave no 
definite clue on the leaf change. 



164 



Flora Malesiana 



[ser. I, vol. 8 2 




The calyx contains water in bud, like other 
coarse-flowered members [Spathodea, Oroxylum, 
etc.). 

Uses. Gamble (Manual Ind. Timb. ed. 1922, 
517) said it is a good timber with close-grained 
wood. Parkinson I.e. found it common in the 
Andaman Is. where it is used for canoes, planking, 
and boat-building; timber smells like teak and 
seems to withstand attacks of white ants. Kurz 
(1870) recorded the largest tree with a trunk of 
2 m 0. In the Natuna Is. it is estimated for building 
boats. 

In the latter islands a decoction of leaves is used 
against fever; in Malaya a hot fermentation is 
applied on the body for stomach disorders 
(Burkill, Diet. 1935, 1623). 

Vern. Bekak gunong, bongli, kaju bonglai, M, 
Malaya, kaju semua, Natuna Is., abeueng laut, 
Meulaboh, Atjeh. 

Note. For obscure reasons Beddome (I.e.) found 
this anomalous in the family, its flower reminded 
of Jasmineae; but he also included Schrebera 
(Oleaceae)\ 



Fig. 29. Tree of Pajanelia longifolia (Willd.) K.Sch. 

showing the sparingly branched, ± pachycaul 

habit ; behind is a limestone hill with dry evergreen 

forest (photogr. Corner, 1935, Sg. Sedili). 



1977] 



Bignoniaceae (van Steenis) 



165 



13. NEOSEPICAEA 



Diels, Bot. Jahrb. 57 (1922) 500, f. 1; Steen. Thesis (1927) 899; Bull. Jard. Bot. 
Btzg III, 10 (1928) 216; Nova Guinea n.s. 8 (1957) 173; ibid. Bot. n. 3 (1960) 15. — 
Haussmannia F.v.M. Fragm. 4 (1864) 148, non Hausmannia Dunker, 1846; K.Sch. 
in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 223; Steen. Thesis (1927) 901. — Nyctocalos 
subg. Haussmannia Seem. J. Bot. 8 (1870) 149. — Pandorea sect. Leptophyllae 
Steen. Thesis (1927), 841, in claw; Bull. Jard. Bot. Btzg III, 10 (1928) 200. —Pan- 
dorea sect. Grandiflores Steen. Nova Guinea 14 (1927) 301, in clav., pro parte. — 
Tecomanthe sect. Aurantiacae Steen. Thesis (1927) 872, in clav.; Bull. Jard. Bot. 
Btzg III, 10 (1928) 203. — Haussmannianthes Steen. Proc. R. Soc. Queensl. 41 
(1929)50. — Fig. 30, 37c. 

Large lianas. Twigs with a distinct gland-field on the nodes. Leaves digitately 
compound; leaflets 3-5, sessile or short-stalked, only articulated at the base of the 
petiolule, mostly unequal, terminal one largest, often with a metallic hue above 
(s.s.), both faces with numerous microscopical glandlets, underneath besides with 




Fig. 30. NeosepU ara van i>utr\ Dii i s. a. Hahit, ' 2 , b. IS of flower, showing also disk and staminode, 

> 2, c .( Sofovanr, 12. N.I*ptophylla(B\ I Steen. d. Just opened capsule tnd one valve from inside, 

„ e. two seeds, - ' j, f. Bower, • ' , ui , 1 bdumann ( >k<w, afta Ddls, </ <■ Bbccari im* 687,/ 

D. Bergmann 261). 



166 



Flora Malesiana 



[ser. I, vol. 8 2 



few, flat, scattered, larger, round glands V4-V2 mm • Thyrses cv>flowered, ter- 
minal, axillary to ramiflorous. Calyx proportionally small, cupular to campanulate, 
truncate and minutely toothed, or shallowly 5-lobed by tearing. Corolla short- 
campanulate or more often ± curved, narrow trumpet-shaped, glabrous, or out- 
side at least the lobes puberulous-papillose; lobes deltoid, valvate, tomentose on the 
inner margin, lobes or tube inside sometimes with larger hairs; tube with a dense 
ring of long hairs at the insertion of the stamens. Stamens 4, didynamous, exserted, 
5th rudimentary; anther-cells divaricate. Disk cupular, enveloping the ovary base. 
Ovary glabrous; style exserted; stigma 2-lamellate, 2-celled, with 2 placentas in 
each cell and 00 ovules. Capsules stipitate, c. 10-20 cm long, narrow oblong, terete 
to broad-ellipsoid in section, beaked; valves boat-shaped. Seeds 00, thin-winged, 
rectangular. 

Distr. Queensland (1 sp.) and Malesia: New Guinea (3 spp.). Fig. 31. 

Ecol. Rain-forests, from the lowland up to c. 2000 m. 

Note. In the Campsis-aUlance possibly closest related to Pandorea, different by the almost regular 
flowers, the valvate corolla lobes, the exserted stamens and style, digitate leaves, and cupular disk clasping 
the base of the ovary. 

KEY TO THE SPECIES 

1. Corolla brown, basal tube, upper tube and lobes all about 1 / 2 cm long; lobes inside bearded, tube on 

one side so 1. N. viticoides 

1. Corolla red or orange, at least 3 cm long, the tube many times as long as the lobes. 

2. Corolla orange, 10 cm long incl. the lobes 3. N. aurantiaca 

2. Corolla red or pink, 2 l / 2 -l cm long incl. the lobes. 
3. Corolla 5-7 cm long, puberulous to glabrous, lobes 1-2 cm long, outside sometimes with dark glands, 
inside sometimes long-hairy. Thyrses axillary. Capsule 13-20 by 3 cm, with woody valves. Seeds 

(incl. wings) 5 by P^cm 2. N. leptophylla 

3. Corolla 2V 2 -3 cm long, glabrous outside except occasionally the puberulous apex of the glandless 
lobes V2 cm long, inside glabrous. Thyrses terminal and from the uppermost leaf-axils. Capsule 
10-15 by 2V2 cm, the valves thin-coriaceous. Seeds (incl. wings) 3 by 1 cm. Queensland. Cf. Steen. 
Nova Guinea n.s. 8 (1957) 174 N. jucunda (F.v.M.)Steen. 



1. Neosepicaea viticoides Diels, Bot. Jahrb. 57 
(1922) 500, f. 1 ; S. Moore, J. Bot. 61 (1923) Suppl. 
38; Steen. Thesis (1927) 900; Bull. Jard. Bot. 
Btzg III, 10 (1928) 217; J. Am. Arb. 28 (1947) 423; 
Nova Guinea n.s. 8 (1957) 174; ibid. Bot. n. 3 
(1960) 15. — Fig. 30a-c. 

High-climbing canopy liana. Leaflets (3-)4-5, 
oblong-elliptic, acuminate, chartaceous, often 
with a metallic hue above, sessile to IV2 cm stalked 
by the tapering base; 8-17 by 3-7V2cm; petiole 
3-10 cm. Thyrses terminal and in uppermost leaf- 
axils, 5-20 cm long. Pedicels c. 1 / 4 - 1 / 2 cm. Calyx 
cupular, 3-5 by 3-5 mm, shallow-lobed, glandless. 
Mature corolla 2'/2~3 cm, outside papillose, basal 
tube about as long as the widened upper tube and 
as the lobes, brown, with dark streaks inside ; lobes 
long-hairy within, 3-4 mm; tube with a long-hairy 
zone inside to the insertion of the stamens. 

Distr. Malesia: East New Guinea (18 collec- 
tions), in W. one. 

Ecol. Rain-forest, not rare, from sea-level to 
c. 1500 m. Fl. Jan.-Oct. 

Notes. A homogeneous species, but the flower 
colour is variously defined probably in part due to 
the age of the flowers : dull ochre, dull yellow with 
maroon markings, brownish olive, dull red or 
brick-purple, brown and purple within; petals 
velvet red. Young leaves are purple. 



No fruit has as yet been collected, and it has only 
once been found in West New Guinea. 




Fig. 31. Range of the genus Neosepicaea Diels; in 

New Guinea 2 (?3) and in Queensland 1 endemic 

species. 



2. Neosepicaea leptophylla (Bl.) Steen. Nova 
Guinea, Bot. n. 3 (1960) 15. — Tecoma leptophylla 
Bl. Rumphia 4 (1849) 35, quoad flor.; Mus. Bot. 1 
(1849) 27; Miq. Fl. Ind. Bat. 2 (1858) 758; Ann. 
Mus. Bot. Lugd.-Bat. 1 (1864) 197; K.Sch. in 



1977] 



Bignoniaceae (van Steenis) 



167 



K.Sch. & Laut. Fl. Schutzgeb. (1900) 540. — 
Gelseminum leptophyllum O.K. Rev. Gen. PI. 2 
(1891) 480. — Pandorea leptophylla Boerl. Handl. 
2 (1899) 600; Diels, Bot. Jahrb. 57 (1922) 449; 
Steen. Nova Guinea 14 (1927) 301, t. 33; Thesis 
(1927) 843 ; Bull. Jard. Bot. Btzg III, 10 (1928) 200; 
Proc. R. Soc. Queensl. 41 (1929) 46, 56. — N. 
superba Steen. Nova Guinea n.s. 8 (1957) 173; 
ibid. Bot. n. 3 (1960) 15. — Fig. 30d-f. 

Large liana, up to 20 m, 3 cm 0. Leaflets 3-5, 
oblong elliptic, acuminate, chartaceous to coria- 
ceous, often with a metallic hue above, base 
cuneate in degree, from sessile to not rarely 
narrowed to a pseudo-petiolule to 3 cm long; 
6-18 by 3-11 cm; petiole 5-12 cm. Thyrses 
obviously axillary, or on old wood, 5-30 cm long. 
Pedicels ' r l' 4 cm. Calyx cupular, not appressed 
to corolla, 4-7 by 6-7 mm, margin ± entire to 
shallowly lobed, glabrous, often with a few glands. 
Mature corolla 5-7 cm (incl. the lobes c. 1-2 by 
x li- i U cm), tube 2\' 2 cm wide at mouth, slightly 
curved, outside glabrous or papillose (except at 
base), purplish (see notes), lobes outside papillose 
or glabrous, with few to several dark glands or 
glandless, inside sometimes with long hairs; tube 
inside glabrous or with a few or a line of long hairs. 
Capsule 13-20 by 2V 2 cm, terete or oval in section, 
the valves woody. Seeds (incl. the wings) 4-5 by 
l 1 2 cm. 

Distr. Malesia: New Guinea. 

Ecol. Rain-forest from the lowland to c. 2000 m. 
Fl. March-May, July-Nov. 

Vern. Ie-up, Kebar (once noted). 

Notes. Fruits have only once been collected by 
Beccari (Arfak Mts). 

All inflorescences hitherto observed are axillary, 
a single one is from old wood. 

The flower colour seems to vary, and to change. 
Vink (BW 11404) noted buds: calyx green, tube 
pale green, lobes purple; submature flower base of 
tube yellow, further orange-red, inside yellow, 
orange-red veined. Pullen (7729) noted on mature 
flowers: pale purple, lobes recurved, inside white 
with purple streaks. Others say merely flowers pale 



violet; Pleyte (887) noted yellow. As in N.jucunda 
the lobes obviously are reflexed in mature flowers. 

I have reduced here N. superba which was dis- 
tinguished by the papillose-puberulous corolla, 
dark glands on the outside of the lobes, and long 
hairs within extending in a narrow line into the 
tube. Three collections have this well expressed 
(Brass 23829, Paymans 53 and Pullen 7729). 
However, there are other specimens defeating these 
characters: Satake 834 and Pleyte 887 have no 
hairs in the tube and hardly any glands on the lobes ; 
Beccari s.n. has a puberulous corolla, but no long 
hairs; Bergmann 226 has an occasional hair in the 
tube and an occasional gland on the lobes, but a 
glabrous corolla; Bergmann 261 has the long hairs 
inside, but a glabrous corolla without glands and 
thus comes nearest to Blume's type. 

The Papuan species is close to N. jucunda from 
Queensland. This has a much smaller, glabrous 
corolla except occasionally the puberulous tip of 
the lobes, the latter constantly much smaller; also 
the thyrses are generally terminal whereas they 
always seem lateral in N. leptophylla; its fruit is 
smaller and the valves much thinner. 



3. Neosepicaea aurantiaca (Diels) Steen. Blumea 
15 (1967) 298. — Tecomanthe aurantiaca Diels, 
Bot. Jahrb. 57 (1922) 497; Steen. Nova Guinea 14 
(1927) 296; Thesis (1927) 874, 834, f. 3d; Bull. 
Jard. Bot. Btzg III, 10 (1928) 204. — Fig. 37c. 

Leaflets 3, coriaceous; petiole 6-7 cm; blade 
oblong, acuminate, cuneate at base, 10—15 by 
4'/2-6 cm; nerves 8-10 pairs. Thyrses axillary; 
peduncle 8 cm. Calyx 5-8 by 8-9 mm. Corolla 
orange, c. 10cm long, 2V4-2 3 / 4 cm wide at apex; 
lobes inside puberulous, IV2-2V2 cm. 

Distr. Malesia: East New Guinea (Etappenberg, 
Sepik Distr. : Ledermann 9561, Bf), once found at 
850 m. 

Note. The cupular calyx and thyrsoid inflores- 
cence stamp this as a Neosepicaea. It might turn 
out to be an exceptionally large-flowered form of 
A', leptophylla. 



14. TECOMANTHE 



Baill. Hist. PI. 10 (1891) 41 ; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 230; 
Boerl. Handl. 2 (1899) 590; Diels, Bot. Jahrb. 57 (1922) 496; Steen. Nova 
Guinea 14(1927) 294; Thesis (1927) 864; Bull. Jard. Bot. Btzg III, 10(1928)201, 
incl. sect. Dendrophilae, Voluhiles el Saxosae Steen. I.e. 205, 208, 210; Pac. PI. 
Areas 1 (1963) 288, map. — Campana Rumph. ex Post & O.K. Lexicon (1904) 95, 
nom. inval. — Pandorea seel. Grandi/hres Steen. Nova Guinea 14 (1927) 301, pro 
parte. — Fig. 32-36. 

Small to large lianas, climbing or creeping (in mountain heaths). Glands on 
twig-nodes small. Leaves 1 -pinnate, 1-7-jugate; leaflets entire or toothed, under- 
neath very finely punctate-glandular. Racemes short, pendent, from efoliate nodes 
on old wood, very rarely axillary or terminal; peduncle short, with some crowded, 
small, sterile bracts at base, in fruit thickening like a brachyblast; rachis short (up 
to c. 7( 13) cm); flowers opposite, in the axil of a small, narrow fugacious bract. 



68 



Flora Malesiana 



[ser. I, vol. 8 2 




1977] Bignoniaceae (van Steenis) 169 

Pedicels with 2 small, narrow bracteoles. Calyx closed in bud, persistent, c. V/ 2 -4 
cm, with 5 fairly large, deltoid ± equal lobes short-hairy along the margin, 
rarely split on one side. Corolla infundibuliform, the basal tube gradually widening 
upwards, ± straight, mostly pink, c. 5-12 cm long, inside near the insertion of the 
stamens stuppose or lax-hairy and sometimes with capitate-glandular papillae; 
limb mostly slightly zygomorphous, very rarely distinctly zygomorphous ; lobes in 
bud narrowly imbricate, mostly deltoid. Stamens didynamous, mostly included, 
5th rudimentary; anther-cells almost free, (in Mai.) c. 4 mm and almost always 
divaricate. Disk thick, annular. Ovary glabrous, in each of the 2 cells with oo rows 
of ovules attached on 2 placentas on the dissepiment; stigma long, filiform, with 2 
spathulate stigmas. Capsule linear-terete or flattened, stipitate and beaked, with 
2 coriaceous or almost woody, smooth, wide or very narrow boat-shaped valves. 
Seeds co, orbicular, with fairly large, thin-membranous wings. 

D i s t r. Species 5, 1 in the Three Kings Is. at the N. tip of New Zealand, 1 in East Queensland, the others 

in Malesia: Moluccas (Ternate, Halmaheira, Ambon, Ceram, Aru Is.), throughout New Guinea (incl. 

Misool, Biak, Jappen, New Britain and Woodlark I., Trobriands), and ?Solomons (Bougainville). Fig. 34. 

Ecol. Primary and secondary rain-forest, mossy forest and mountain heaths, from sea-level up to 

3100m. 

One collector noted honey in the flowers. It is likely that the diurnal flowers are bird-pollinated, but 
there are no records. 

Tecomanthe speciosa was derived from cuttings of the single plant found in nature. It has in cultivation 
produced flowers and can obviously propagate by self-pollination. The corolla is sometimes (in the 
herbarium) already ± open before full maturity. In one case rather long, unbranched, pendent, aerial 
roots were observed emitted from a node. In juvenile specimens leaves tend to be more toothed than in 
mature foliage. 

Tax on. As I stated before Tecomanthe belongs with Pandorea to a distinct circum-Pacific affinity of 
lianas, including Campsis in East Asia and North America and Campsidium in Chile, all sharing a similar 
shape of flower and fruit. The South African genus Podranea is more remote and with its inflated calyx 
and linear capsules possibly more allied to Tecoma. The four genera can be keyed out as follows: 
1. Flowers in racemes, opposite. Evergreen, not climbing with roots. Calyx with well-developed lobes, 
1-4 cm long. Corolla with a hair-ring (lax or stuppose, sometimes replaced in part by capitate- 
glandular papillae) near the insertion of the stamens. 
2. Racemes almost always on the old wood, rarely axillary or terminal. Peduncle at the base with 
crowded sterile bracts, rachis up to 7 cm, pendent, flowers close together. Calyx large, 1 l / a -4 cm. 
Corolla 6-12 cm, tube not contracted below the slightly or distinctly zygomorphous limb; lobes 
deltoid, in bud narrowly imbricate. Anthers mostly included, cells almost always divaricate. Valves 
of the capsule widely or narrowly boat-shaped ; endocarp not removable Tecomanthe 

2. Racemes terminal on leafy twigs. Peduncle at base without bracts; flower pairs spaced. Calyx 
c. 1 cm. Corolla c. 3'/ 2 cm long, tube contracted below the limb; lobes rounded. Stamens ± exserted; 
anther-cells free but parallel. Capsule narrowly elliptic-oblong, with removable papery endocarp. 
Chile Campsidium 

1. Flowers paniculate in terminal thyrses, very rarely depauperate in racemes but then the calyx much 
smaller than 1 cm. Corolla lobes widely imbricating. Anther-cells divaricate. 

3. Deciduous, climbing with roots. Leaflets 5 pairs, distinctly sharply serrate. Calyx large, c. 17 2 -3 cm, 
with large lobes. Corolla inside glabrous, large, c. 5 l l 2 -% cm Campsis 

3. Evergreen, not climbing with roots, rarely erect (in arid country). Leaflets 1-7 pairs, not distinctly 
sharply serrate in mature specimens. Calyx small, stunted or short-lobed, 2V2-8 mm. Corolla smaller, 
I ' . 3'/ 2 ( 5) cm, almost always with a hair-ring near the insertion of the stamens and long hairs one- 
sided in throat and upper part of tube Pandorea 

Uses. Almost all species are ornamental and can be cultivated in tropical and subtropical countries; 
they are not hardy. Propagation by cuttings or seed. 

KEY TO THE SPECIES AND SUBSPECIES 

I. Corolla cream-coloured, woolly tomentose in ihc upper half, <"> 8 cm long, the limb very zygomorphic. 
Stamens exscrted, anthers c. 10 mm, with parallel cells. Calyx often split on one side to the base, the 

Fig. 32. Temnuinihr dciulmplulu (\\\ ) K.S< 11 .1 Habit, ' , h. pistil, c. anther, both cnlai fed, d. capsule, 

/ tematttuU Btun. e. 1 cat, 1 g. inflorescence in bud and Bower, all ' . h < Sol ovary (a e 

after Mi 1 mi , ,1 I A I 58636, t h Mm, din 1201). 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 33. Tecomanthe dendrophila (Bl.) K.Sch. in the mossy forest on Mt Cycloop, N. New Guinea, at 

1200 m (photogr. Van Royen). 



1977] Bignoniaceae (van Steenis) 171 

lobes very unequal. Leaflets 5, orbicular-elliptic, apex broadly rounded to slightly notched, 8-18 by 
5-11 cm, fleshy, coriaceous when dry. Capsule terete, pointed at both ends, c. 16 by 2V2cm; valves 
thick, almost woody. Seeds 3-4 by 1\\ cm (incl. wings). Cf. Hunter, Rec.Auckl. Inst. Mus.5 (1958)41, 
pi. 6-7; Hunt, Bot. Mag. 179 (1972) t. 618. Three Kings Is. (New Zealand) T. speciosa Oliv. 
1. Corolla at most puberulous in the upper half of the lobes, not very zygomorphic. Anther-cells 3-4 mm 
long, divaricate. Calyx never split on one side to the base, lobes equal or unequal. Leaflets acute to 
acuminate. 
2. Uniseriate hairs near the insertion of the stamens very lax, few or almost absent. Rachis of raceme 
glabrous. Corolla 4\ 2 -7 cm long. 
3. Leaflets in 3-4 pairs, c. 3'/ 2 -6 by 1V2-3 cm, about twice as long as wide, herbaceous, veins between 
the main nerves usually distinct; lower lateral petiolules 0-3 mm. Calyx c. l l / 3 cm long (incl. lobes). 
Corolla whitish, later pink tinged, 4V 2 -6cm, the lobes 2-5 by 10-15 mm, with dark dots in trans- 
parent view. Stamens as long as the style, ± exserted. No capitate-glandular papillae near the inser- 
tions of the stamens 2. T. ternatensis 

3. Leaflets in 2 pairs (4 or 5), c. 3-$ l l 2 by lV4-4cm, at least twice as long as wide, often narrower, 
obviously rather fleshy, at base rounded to truncate, often oblique, veins between the 4-5 main 
nerves hardly visible; lower lateral petiolules 4-9 mm, longer than upper ones. Calyx 2 l / 2 -3 cm long 
(incl. lobes). Corolla tube pale, limb pink to rosy-purplish, tube marked with purplish lines inside, 
4 1 j-7 cm, lobes ovate-triangular, ( 3 / 4 -)l-l 3 / 4 by ( 3 / 4 -)lV 2 -l 3 / 4 cm, without dark dots. Stamens ± 
shorter than the style, not exserted. Near the insertions of the stamens mainly capitate-glandular 
papillae and no or few uniseriate hairs. Capsule 5\/ 2 by 2 cm. Cf. C. T. White, Queensl. Nat. 4 (1920) 
100, f. ; Steen. Proc. R. Soc. Queensl. 41 (1929) 49. Queensland T. hillii (F.v.M.) Steen. 

2. Hairs near the insertion of the stamens in a stuppose ring. Corolla 6-13 cm. Rachis of inflorescence 
often puberulous. 

4. Leaves 1-2-jugate; leaflets fairly large, averagely 5-10 by 2 l l 2 -5 cm, mostly herbaceous, usually 
entire, occasionally with a few coarse teeth to apex; rachis not winged. Lateral pedicels 2-8 mm. 
Racemes finally rich-flowered (6-20), on the old wood of coarse lianas. Capsule broad-elliptic in 
section, c. 17-22(-30)cm long, c. 3(-3 3 / 4 ) cm wide and thick, the valves wide-boat-shaped, hard, 
almost woody. Seeds including wing 2 l / 2 -3 l /2 by l'/ 4 -l '/2 cm 1. T. dendrophila 

4. Leaves 2-7-jugate; leaflets small, averagely l'/2-4(-8) by l l 2 -l 3 U(-3 l l 2 ) cm, herbaceous to coriaceous, 
the margin almost always toothed ; the sulcate rachis (very) narrow-winged ; lateral pedicels 0-2 mm. 
Racemes pauciflorous (2-6), lateral or terminal, on small, slender lianas. Capsule flat and com- 
pressed, 8-14 cm long, the valves very much compressed-boat-shaped, coriaceous. Seeds including 
wings c. IV2-2 by 1-172 cm. 
5. Leaves 3-6(-7)-jugate, 4-jugate leaves always present. Leaflets thick-coriaceous to chartaceous 
(or even herbaceous), elliptic with usually short, acute apex, brittle in the herbarium, the nerves 
and midrib usually impressed above, never strongly prominent. Calyx IV2-3 cm. Corolla 6-10 cm. 

Staminode usually less than 1 cm long 3. T. volubilis 

5. Leaves 2(-3)-jugate, 4- and more-jugate leaves absent. Leaf-apex acute to cuspidate. Staminode 

lV2-3cm. 
6. Leaflets coriaceous, tough when mature, not easily breakable, sharply toothed, with nerves and 
veins conspicuously prominent on both sides. Calyx (l 3 / 4 -)2-3V 2 cm. Corolla 7-12 cm. 

3b. T. volubilis ssp. tenax 

6. Leaves coriaceous to herbaceous, brittle in the herbarium, toothed but not sharply so; midrib 

and veins flat or slightly sulcate above, somewhat prominent beneath. Calyx IV2-2V2 cm. Corolla 

6-8 cm 3a. T. volubilis ssp. silvicola 

1. Tecomanthe dendrophila (Bl.) K.Sch. in K.Sch. 373. — Campsis amboinensis Seem. I.e. 374. — T. 

& Laut. Fl. Schutzgcb. (1900) 539; Rech. Denk- bureavii Baill. Hist. PI. 10 (1891) 41; K.Sch. in 

schr. K. Ak. Wiss. M.-N. Kl. Wien 89 (1913) 603; E. & P. Nat. Pfl. Fam. 4, 3b (1894) 230. — Gelse- 

Diels, Bot. Jahrb. 57 (1922) 496; Steen. Nova minum amboinense et dendrophilum O.K. Rev. Gen. 

Guinea 14 (1927) 297; Thesis (1927) 880; Bull. PI. 2 (1891) 479. — Pandorea dendrophila Boerl. 

Jard. Bot. Btzg III, 10 (1928) 206; Laut. Bot. Handl. 2 (1899) 600. — Pandorea amboinensis 

Jahrb. 62 (1928) 292; Loihian, J. R. Hort. Soc. 83 Boerl. I.e. — T. gloriosa S. Moore, J. Bot. 61 

(1958) 295; Sykis, Stud. Cult. PI. N.Z. 1 (1966)43, (1923) Suppl. 38; Shin. Nova Guinea 14(1927) 

f. 18; Hi hkiois. I I. Trop. Climb. (1976) 73, f. 97. 299; Thesis (1927) 888; Bull. Jard. Bot. Btzg III. 

— Campana rubra Rumph. Herb. Amb. (1755) 10 (1928) 210. - T. venusta S. Moore, J. Bot. 61 

Auct. 42; an Pandorea' 1 , Mikk. Int. Rumph. Herb. (1923) Suppl. 38; Steen. Nova Guinea 14 (1927) 

Amb. (1917) 469. — Tecoma dendrophila Mi. 298; Thesis (1927) 897, f. 5b, incl. var. parvijiora 

Ri mi'hia 4(1849) 35, et Dendrophila trifoliate Bl. Shin.; Bull. Jard. Bot. Ht/g III. 10 (1928) 216; 

sub. t. 190; Mus. Hot. I (1849) 25; M10. II. Ind. Hum, Hot. Man. 1X0 (l ( >75) I. 693; Hi km ms, Fl. 

: HX58) 757; Ann. Mus. Bot Lugd Bat. I [top. Climb. (1976) 71, f. 96, col. dL 7. — T. elltp- 

(IXM) 197, k.s< 11. Bot. Jahrb. 9 (ixx7» 218; Fl. tlea Shin. Nova Guinea 1 » (1927) 296, 1. 3 41); 

Wilh I ,nul(IXX9) 123; Warm. Hoi. Jahrb. 13 Ihcsis (1927) 876, f. 5a; Hull. Jard. Hot. Htzg III, 

(1X91)418; l-.v.M. Deecr. Not. 9 (1890) 10(1928)205. r. acutifolla Stebn. Nova Guinea 

Tecoma ambolnensU BL. Kumphia 4 (1X49) 15; 14(1927)297; Hk.i-. (1927) 879; Hull. Jard. Hoi. 

H..t. I (1849)26; Miq. Fl. Ind. Bat 2 (1858) Bt/g ill. 10(1 r. amboinensis Stbbn. 

757. - Campus dendrophila Sum J Bot 5 (1X67) Nova Guinea 14 (1927) 29X; Thesis (1927) 890; 



172 



Flora Malesiana 



[ser. I, vol. 8 2 



Bull. Jard. Bot. Btzg III, 10 (1928) 211. — T. 
gjellerupii Steen. Nova Guinea 14 (1927) 298; 
Thesis (1927) 896; Bull. Jard. Bot. Btzg III, 10 
(1928) 215. — Fig. 32a-d, 33. 

A tall liana, up to 20(-30) m. Leaves 1-2-jugate; 
leaflets ovate to elliptic or oblong-lanceolate, 
herbaceous to chartaceous, entire or with a few 
coarse teeth to the top, apex rather blunt to acumi- 
nate, (3-)5-l 3 by ( 1 l l 2 -)2 l l 2 -l cm ; nerves flat above 
or slightly impressed, prominent beneath; rachis 
not winged; lateral petiolules 2-8 mm. Racemes on 
the old wood, the rachis c. l l 2 -l(-\3) cm, with 
usually 6-20 densely set flowers; pedicels 1-2 cm. 
Calyx herbaceous to ± coriaceous, (1 74-) 1 1 / 2 -4 cm 
long, for V4-V2 incised, greenish tinged red to 
purple-brown, the lobes triangular, blunt to 
cuspidate, 8-15 by 5-10 mm, midrib prominent or 
not. Corolla 7-11 cm long including the broad 
triangular acutish to blunt lobes i U~l 1 l 2 by 1-2 cm, 
the tube pink, rosa or pale carmine, the lobes 
creamy to yellowish, sometimes streaked with 
purple lines, or pink all over, inside near the inser- 
tion of the stamens stuppose-hairy. Anthers c. 

4 mm long, divaricate. Capsule almost cylindric, 
stiped and beaked, 17-22(-30) by 3-3 3 / 4 by 3 cm, 
with hard, almost woody, boat-shaped valves. 
Seeds including the thin wing 2 1 / ' 2 -3'/ ' 2 by IV4- 
IV 2 cm. 

Distr. Malesia: Moluccas (Ambon, Ceram, Aru 
Is.), New Guinea (throughout, and incl. Misool, 
Jappen, Biak, Woodlark I. & New Britain), and 
?Solomons (Bougainville), 120 collections. 

Millar & Vandenberg collected this species 
(NGF 48505) at Arawa Plantation, Kieta Sub- 
distr., Bougainville, cultivated in a garden 'from a 
native vine'. I feel not certain that it is native in 
Bougainville; it might have been introduced from 
Papua. 

Ecol. In swampy or dry rain-forests, sometimes 
riverine forest, once on limestone, from sea-level 
up to c. 1500 m. Fl. April-Dec.,//-. June-Nov. 

Vern. Asee, Maibrat lang., fiyo, Wapi lang., 
Marok; Sepik: gwimbipuk, gwoimbipok, Waskuk, 
sanie, Ambuti, ilei, Wagu, yakomenga, Narak & 
Ganja, Mt Hagen. 

Notes. Through the great increase in collections 
it has appeared impossible to maintain several 
formerly described species. The characters of the 
leaves, calyx and corolla show transient, not 
correlated variation. Though the number of herba- 
rium collections in which 1- and 2-jugate leaves 
occur together is restricted, they do occur on one 
plant in cultivation and in the forest according to 
collectors. The calyx shows a great variation in 
size and degree of incision. Puberulous hairiness 
may occur on the rachis, pedicels, the calyx, the 
midrib and nerves beneath, the apical part of the 
corolla-lobes, and on the leaf-rachis. Aberrations 
are sometimes found in individual specimens: a 
multi-lobed, wide calyx in BW 11242; a very thin, 
tortuous, 20 cm long rachis of a lax raceme in NGF 
11866 and BW 13352; once a leaf with 6 and 7 
leaflets; once long unbranched roots produced 
from a node of the old wood (Janowski 427) ; an 
axillary raceme (Brass 28745). 

Seemingly open flowers measure sometimes only 

5 cm, but I assume this to be caused in drying of 
immature flowers and tardy growth. 




Fig. 34. Range of the genus Tecomanthe Baill. 
Figures above the hyphen indicate endemic species, 
those below the hyphen non-endemic species. The 
Australian T. hillii (F.v.M.) Steen. occupies two 
areas. 



2. Tecomanthe ternatensis Steen. Thesis (1927) 
893 ; Bull. Jard. Bot. Btzg III, 10 (1928) 214, f. 1. — 
Fig. 32e-h. 

Large liana, up to 30 m or more ; stem to arm- 
thick. Leaves 3-4-jugate; leaflets elliptic, herba- 
ceous, base ± rounded, apex short-acute, with 
some teeth towards the apex, c. Vl 2 -1 by IV2-3 cm; 
veins between the nerves usually distinct; lower 
lateral petiolules 0-3 mm. Racemes on the old 
wood, 2-6 cm long, densely rich-flowered. Pedicels 
c. 1 cm. Calyx pale green, 1 1 / 2 -2 cm long, the lobes 
deltoid, 7 by 5 mm. Corolla whitish, later tinged 
pink, 5-6 cm long including lobes, lobes wide and 
short, c. 2-5 by 10-15 mm, with dark spots in 
transparent view; tube near the insertions of the 
stamens with very few hairs and no capitate-glandu- 
lar papillae. Stamens as long as the style, ± 
exserted. 

Distr. Malesia: Moluccas (Ternate, Halma- 
heira) and N W. New Guinea (Biak I.), 5 collections. 

Ecol. Primary and secondary forest, in Biak on 
coralline limestone, in the Moluccas at 500-600 m. 
Fl. Sept.-Nov., April. 

Note. The Biak specimens are rather poor but 
clearly belong to this species. 

3. Tecomanthe volubilis Gibbs, Arfak (1917) 179; 
Diels, Bot. Jahrb. 57 (1922) 498; Steen. Nova 
Guinea 14 (1927) 299; Thesis (1927) 885; Bull. 
Jard. Bot. Btzg III, 10 (1928) 209. — T. nitida 
Steen. Nova Guinea 14 (1927) 299, t. 33; Thesis 
(1927) 887; Bull. Jard. Bot. Btzg III, 10 (1928) 209. 
— T. arfak i Steen. Nova Guinea 14 (1927) 300, 
t. 34B; Thesis (1927) 884, f. 5d, j; Bull. Jard. Bot. 
Btzg III, 10 (1928) 208. — Fig. 35a-c. 

Small, slender climber, 2-5 m. Leaves 3-6(-7)- 
jugate, with 4-jugate leaves always present; 
leaflets mostly glossy on both sides, dark green 
above, pale beneath, mostly coriaceous, brittle in 
the herbarium, ovate, obovate to elliptic, rarely 
lanceolate, base usually cuneate, apex acute, rarely 
blunt, margin in exposed places recurved, usually 



1977] 



Bignoniaceae (van Steenis) 



173 



with 1-5 pairs of bluntish teeth, 3 U-2 l l 2 by l /sr 
lVzCm, usually sessile but lateral pedicels up to 
2 mm, nerves above usually impressed, beneath 
usually prominent, often nigrescent in sicco. 
Racemes axillary or terminal, rachis i l 2 -2 i / 2 cm; 
flowers 1-3 pairs, pedicels ^-l'^cm, both often 




Fig. 35. Tecomanthe volubilis Gibbs. a. Habit, b. 
unripe capsule, c. leaf. — T. volubilis \sp. silvicola 
Srt i •.. d. Leaf. — T. volubilis ssp. tenax Sin M, t. 
Leaf, underside, f. leaflet, upper surface. All X '/i 
(a van Row. (t Si ii mi k 7417, b Bkass 9052, 
c BW 3050. d LfcUERMANN 12904, e f Kai.kman 
5178). 



lax-puberulous. Calyx green suffused with red, 
coriaceous to herbaceous, 1 */2-3 cm, tube as long 
as or up to 2 times as long as the acute triangular 
lobes. Corolla pink to carmine, inside creamy, 
sometimes streaked red inside, 6-8(-10) cm includ- 
ing the triangular, acute lobes Vj 2 -2 l l 2 by l l / 2 - 
3 cm. Staminode usually less than 1 cm. Capsule 
11-14 by 3-4 cm, compressed; valves coriaceous. 
Seeds c. \ l j 2 cm 0. 

Distr. Malesia: New Guinea (Mts Arfak, incl. 
Nettoti, Tamrau; Wissel Lakes, Wichmann, 
Carstensz, Lake Habbema, Star Mts, Telefomin, 
Bosavi), 20 collections. 

Ecol. Mossy thickets and heaths, often burned, 
ridges in high forest, open scrub and secondary 
forest, sometimes on peaty soil or on limestone, 
1250-3100 m. Fl. Sept.-Febr. (once April), fr. 
Aug., Nov., Jan., April. 

Vern. Basengga, Hattam lang., Arfak, daibuda, 
dibibuda, Kapauku lang., Wissel Lakes. 

Note. At lower altitude and more shaded locali- 
ties leaflets tend to be larger and less rigid and 
coriaceous, with less impressed nerves above and 
flat leaf margin. 

3a. ssp. silvicola Steen. nov. ssp. — T. saxosa 
Diels, Bot. Jahrb. 57 (1922) 498; Steen. Nova 
Guinea 14 (1927) 889, f. 5c; Thesis (1927) 297; 
Bull. Jard. Bot. Btzg III, 10 (1928) 211. — T. 
cyclopensis Steen. Nova Guinea 14 (1927) 298, 
t. 34A; Thesis (1927) 895; Bull. Jard. Bot. Btzg III, 
10 (1928) 214. — T. nitida (non Steen.) Herklots, 
Fl. Trop. Climb. (1976) 73, f. 98. — Fig. 35d. 

Differt a T. volubili foliolis 2(-3)-jugatis, apice 
acutis vel cuspidatis; staminodiis plerumque Vj 2 - 
3 cm longis. — Typus: van Royen & Sleumer 7117 
(L), NW. New Guinea, Vogelkop Peninsula, 
Tamrau Mts, 1350 m. 

Slender liana. Leaflets 2(-3) pairs, mostly her- 
baceous to chartaceous, usually elliptic to ovate- 
oblong, or lanceolate-oblong, entire but mostly 
with several pairs of teeth, base cuneate to rounded, 
apex acute, lateral leaflets l 3 / 4 -5(-8) by 3 / 4 -2V 2 (-3) 
cm, terminal one longest ; pedicels 1 l 2 -6 mm. 
Racemes axillary or on old wood. Calyx usually 
herbaceous, 172-3(-3V2) cm, halfway incised or ± 
less. Corolla 5— 7(— 9— 1 1) cm, including the lobes. 
Capsule flat, 8-12 by 2-3 cm. Seeds VI 2 -2 by 
1-172 cm. 

Distr. Malesia: New Guinea (throughout, but 
far more common in East New Guinea, common in 
Morobe Distr.), 30 collections. 

Ecol. In the understorey of Nothofagus- 
Araucaria and Castanopsis forest, more rarely in 
moss forest and on ridges, mostly in fairly tall 
forest, (lOOO-)15OO-24O0(-3OO0) m (once found at 
80 m between Hollandia and Sentani). Fl. Jan.- 
Dec.,/r. March, June, Oct. 

Note. I regard this montane forest plant to 
represent a race of T. volubilis; it is rather variable 
in foliage, some specimens looking transitional to 
low altitude specimens of ssp. volubilis. 

3b. up, tenax Steen. nov. ssp. — Fig. 35e-f. 

I)i flirt a foliolis 2( l)-jugati\, coriaceis, venis 
ntrvtsqtu utrtnqtu perspicut promintntibtu. — 
Typus: LAE 60706, leg. CkOFI ft til.. E. Papua. 
s. slopes of Mt. Giluwe, u 1' s. L43°55' E, fl. fr. 
25 xn 1973 (L, isoin LAE). 



174 



Flora Malesiana 



[ser. I, vol. 8 2 



Smallish liana, 2-10 m; branchlets puberulous 
to subglabrous. Leaflets 2-jugate (very rarely 1- or 
3-jugate), dark green above, pale green beneath, 
glossy on both sides, ovate to lanceolate, coria- 
ceous, not nigrescent, 2-5 l j 2 (-%) by l-2(-4)cm, 
terminal one largest, very tough, margin especially 
towards apex with sharp teeth. Racemes axillary, 
rarely terminal, lax, with (1— )2— 3 pairs of flowers; 
rachis and pedicels ± lax-puberulous; rachis 
1-5 cm; pedicels VU-2 l j 2 cm. Calyx mostly thin, 
2 1 / 4 -3 1 /2cm, tube 1 1 ; 2 -2(— 3) times as long as the 
lobes ; reddish white to green with pink tinge ; lobes 
triangular, mucronate. Corolla pink to crimson 
(once noted white), the lobes yellowish to white, 
streaked red within, 7-12 cm long including the 
triangular lobes c. \ l l 2 by \ l l 2 cm. Capsule 8-14 
by 2-3 cm (septum 2'/ 2 cm stalked). Seeds c. 1 l l 2 
by 1 cm including wing. 

Distr. Malesia: New Guinea: Papua (E., S. & 
W. Highlands, largely c. 6° S and 143-144° E), but 
also in West New Guinea (Bernard Camp, Iden- 
burg R.). Fig. 36. 

Ecol. Mountain forest, often mossy, often with 
Nothofagus dominating, sometimes in shrubs 



bordering glades, 1800-3000 m. Fl. June, Sept- 
Jan. (once in May), fir. (twice) July, Dec. 

Vern. Tserki, Enga lang., taugurapu, Tari, 
igidumbroki, Mendi lang. 

Note. The material is very homogeneous in 25 
collections seen and though fertile characters with 
ssp. silvicola are overlapping it can easily be recog- 
nized vegetatively without any transitions. 











Wi ** 




7 ^ 


jU J 






» 


V__Oo 


. . 


y\ 


x -* - , 



Fig. 36. Localities of Tecomanthe volubilis Gibbs 
ssp. tenax Steen. 



15. PANDOREA 

Spach, Hist. Veg. 9 (1840) 136; Mon. (1864) 49; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 
3b (1894) 230; Steen. Nova Guinea 14 (1927) 301, inch sect. Parviflores Steen., in 
clav.; Thesis (1927) 294; Proc. R. Soc. Queensl. 41 (1929) 43. — Tecoma sect. Pan- 
dorea'E^DL. Gen. PI. (1839) 711 ; DC. Prod. 9 (1845) 225. — Tecomanthe sect. Mon- 
tanae Steen. Bull Jard. Bot. Btzg III, 10 (1928) 204. — Fig. 37, 39. 

Lianas, only exceptionally (in arid countries) erect. Glands on twig-nodes small. 
Leaves (l-)2-4(-7)-jugate; leaflets with microscopical glands and not rarely with 
few to many larger crateriform scattered glands underneath ; petiole in some species 
with some ventral large glands near the base. Thyrses terminal, sometimes addi- 
tional partial axillary thyrses in the upper leaf-axils; peduncle without sterile 
bracts at the base; depauperate thyrses may appear occasionally as racemes. 
Calyx closed in bud, small (less than 7 mm), cup-shaped to campanulate, stunted 
or very shallowly lobed, sometimes tearing. Corolla generally small, the tube 
cylindric or infundibuliform, incl. lobes at most 5 cm long, limb usually zygomor- 
phous, the lobes small or large, in bud widely imbricating; throat and ventral side 
of the tube mostly long-hairy and often with a hair-ring near the insertion of the 
anthers. Stamens didynamous, almost always inserted; anther-cells divaricate, 
c. 172 - 2 mm long; 5th rudimentary. Disk annular or ± pulvinous. Ovary elongate, 
each cell with 2 placentas and many ovules. Capsule stipitate, ± beaked, rather 
thick, terete or ± flattened; valves widely boat-shaped, firmly coriaceous; dis- 
sepiment flat, oblong, thickish, with marginal seed-scars. Seeds many, roundish, 
thin-winged. 



Distr. Six spp., Central, N. & E. Australia, Tasmania, Lord Howe I., New Caledonia, Solomons 
(Bougainville), and East Malesia: New Guinea (New Britain included), Moluccas, and the Lesser Sunda 
Islands (Lombok, Flores, Timor). Fig. 38. 

Ecol. In Malesia in rain-forest, from sea-level to 2450 m. 



1977] 



Bignoniaceae (van Steenis) 



175 



KEY TO THE SPECIES 

1. Corolla tube glabrous outside, usually rather wide, without hair-ring inside near the insertion of the 
stamens. Base of filaments glandular and also the ovary with similar sessile glands. Calyx cupular, 
c. 2-3 mm. Venation not prominent above. Filaments inserted close to the base of the tube, which is 

not narrowed 1. P. pandorana 

1 . Corolla tube (except at the very base) outside puberulous. Base of filaments without sessile glands. 

Venation above mostly prominent. 
2. Flowers large, 4-5 cm long (incl. lobes), white with crimson throat, the tube c. twice as long as the 
rounded lobes. Leaves 2-jugate, the entire leaflets blunt at apex, nerves and veins not distinctly 

prominent above. Queensland and New South Wales P. jasminoides (Lindl.) K.Sch. 

2. Flowers much smaller, at most Vl 2 cm, the tube 4-6 times as long as the lobes; lobes less than c. 1 cm 
diameter. Leaflets acute, nerves and veins usually distinctly prominent above, mostly toothed towards 
the apex. 
3. Corolla 12-15 mm long, narrow-cylindric, the tube c. 6 times as long as the lobes. No hair-ring at 
the staminal base. Calyx 3-5 mm. Petiole above at base with one or a few large glands; leaf-rachis 
narrowly winged. 
4. Leaflets entire, 3-4-jugate, 5-12 by 2-5 cm. Corolla c. 12-13 mm, tube ± curved, c. 2-3 mm wide, 
no beard in the mouth and tube, lobes c. 2 mm. Ovary orbicular, with sessile glands. Venation on 
upper surface of leaflets raised, but not fine-tessellate. Flowers cream-coloured, lobes and throat 
pink-shaded. Queensland. Cf. Steen. Proc. R. Soc. Queensl. 41 (1929) 46, f. 1. 

P. baileyana (Maid. & R. T. Baker) Steen. 

4. Leaflets apically toothed, (l-)2-3-jugate, 6-8V2 by VI 2 -4 cm. Corolla tube straight, c. 20 mm long, 
c. 5 mm wide, mouth and upper part of tube inside bearded, lobes c. 3-4 mm. Ovary obconical- 
oblong, eglandular. Venation on upper surface of leaflets raised, fine-tessellate. (Corolla tube 
yellow, the lobes white) 2. P. stenantha 

3. Corolla 20-35 mm long, the tube c. 4 times as long as the lobes, the mouth and upper part of the 
tube inside bearded. Calyx c. (>-l l j 2 mm. Petiole without glands; rachis narrowly winged; leaflets 
dentate in upper part, venation not fine-tessellate raised above. 

5. Corolla straight, tubular, 20-25 mm long, with yellow tube, the lobes white to pale red or streaked 
red, inside with a distinct hair-ring near the insertion of the stamens. Ovary ± conical, eglandular. 
Pedicels slender, 1-2 cm 3. P. montana 

5. Corolla c. 30-35 mm long, white with pale yellow mouth, infundibuliform, with a fairly narrow 
lower part of the tube, widened apically, inside at the insertion of the stamens with a few hairs. 
Ovary ellipsoid, with sessile glands. Pedicels c. 5 mm. Queensland. Cf. Steen. J. Arn. Arb. 12 
(1931) 149, pi. 35 P. nervosa Steen. 



1. Pandorea pandorana (Andr.) Steen. Bull. Jard. 
Bull. Btzg III, 10(1928) 198; Proc. R. Soc. Queensl. 
41 (1929) 43; J. H. Willis, Handb. PI. Vict. 2 
(1972) 578; Beadle, Evans & Carolin, Fl. 
Sydney Reg. ed. 2 (1972) 502; Herklots, Fl. Trop. 
Climb. (1976) 69, f. 91. — Bignonia pandorana 
Andr. Bot. Rep. 2 (1800) t. 86. — Bignonia 
pandorea Vent. Jard. Malm. (1803) t. 43. — 
Bignonia pandorae Sims, Bot. Mag. 22 (1805) t. 865. 
— Tecoma australis R.Br. Prod. (1810) 471 ; DC. 
Prod. 9 (1845) 225, incl. var. meonantha (Link) 
DC; Bth. Fl. Austr. 4 (1869) 537; Bailey, 
Queensl. Fl. 4 (1901) 1134, incl. var. pandorea 
(Vent.) Bailey (= var. typ.), var. meonantha ei 
var. linearis Bailey, I.e. pi. 45. — Bignonia australis 
Ait. Hort. Kew. ed. 2, 4 (1814) 34. — Bignonia 
meonantha Link, En. Berol. 2 (1822) 130. — 
Tecoma meonantha Sweet, Hort. Brit. (1827) 284; 
G. Don, Syst. 4 (1838) 224; Harris, Wild Fl. Austr. 
(1938) 151, pi. 6 (as T. australis). — Tecoma 
diver sifoliaG. Don, Syst.4 (1838) 225; DC.Prod.9 
(1845) 225. — P. australis S\>\< n. Hist. Nat. Veg. 9 
(1840) 136; K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b 
(1894) 230; Dim*, Bot. Jahrb. 57 (1922) 498; 
Sue. Nova Guinea 14 (1927) 302; Thesis 
(1927) 859, in<l. ssp. pandorea Srn h. i.e. B61, up. 

meonantha Siihv I.e. 862, cl ssp. linearis S11 1 •■ / < 

163; Down BibL Bot. 22 (1929) 1153, incl. var. 

oxleyi, nom illeg., ct var. meonantha. I.e. I 154. 
Tecoma florihunda < < NX t\ IX I'rod. 9 (1845) 
225. Tecoma oxleyi Cunn. ex DC. Lcj J. M. 
Hia'k, Trans. R. Soc. S. Austr. 39 (1 ( >15) 836; 



White & Francis, Proc. R. Soc. Queensl. 37 
(1926) 166; Harris, Wild Fl. Austr. (1938) 151, 
pi. 39. — Tecoma ochroxantha Kth & BoucHE,Ind. 
Sem. Hort. Berol. (1847) 12, sec. Bth. 1869. — 
Tecoma leptophylla Bl. Rumphia 4 (1849) 35; 
Steen. Nova Guinea 14(1927) 301, t. 33, pro parte, 
pro fol. sol. — Tecoma austro-caledonica Bureau, 
Bull. Soc. Bot. Fr. 9 (1862) 163; Maid. Proc. Linn. 
Soc. N.S.W. 39 (1914) 382; Compton, J. Linn. 
Soc. Bot. 45 (1921) 373. — Tecoma ceramensis 
T. & B. Nat. Tijd. N. I. 25 (1863) 412; Miq. Ann. 
Mus. Bot. Lugd.-Bat. 1 (1864) 197, t. 5, incl. var. 
elliptica Miq. — P. austro-caledonica Seem. Gard. 
Chron. (1870) 1085; Baill. Hist. PI. 10 (1891) 40; 
K.Sch. in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 230; 
Guillaumin, Fl. Nouv. Cal. (1948) 317 ('austro- 
caledonicum); Heine, Fl. Nouv.-Caled. 7 (1976) 
87, pi. 20. — Campsidium filicifolium Bull, 
Wholesale List New, Beaut. & Rare PI. (1874) fig. ; 
Cat., ex Johnson & Hogg, J. Hort. 51 (1874) 366; 
A. van Geert, Cat. n. 74 (1874); T. Moore, Fl. & 
Pom. ( 1 874) 280. — Tecoma filicifolium Nicholson, 
Diet. Gard. 4 (1887) 13; cf. Steen. Blumea 15 
(1967) 146. — Gelseminurn pandorea et ochroxan- 
thum O.K. Rev. Gen. PI. 2 (1891) 480. — P. cera- 
mensis Maim. Mist. PI. 10 (1K9I) 40; K.Sch. in 
E. & P. Nat. Pfl. Fam. 4, 3b (1 894) 230 ('ceramic a'); 
Smin. Nova Guinea 14 (1927) 302; Thesis (1927) 
X52. lemma pandorana Skills, U.S. Dep. 

Agric. Bur. PI. Ind. Bull. 282 (1913) 62. — P. 
aeutifolui Shin. Nova Guinea 14 (1927) 303, 
t. 34C; Thesis (1927) 855; Bull. Jard. Bot. Btzg III, 



176 



Flora Malesiana 



[ser. I, vol. 8 2 




c Ledermann 9561). ' 



1977] 



Bignoniaceae (van Steenis) 



177 



10 (1928) 196. — P. poincillantha Steen. Nova 
Guinea 14 (1927) 302; Thesis (1927) 857, incl. var. 
fragrans Steen. — Tecoma doratoxylon J. M. 
Black, Trans. Proc. R. Soc. S. Austr. 51 (1927) 
383; Gardner, En. PI. Austr. Occ. (1930) 118. — 
P. doratoxylon J. M. Black, Trans. Proc. R. Soc. 
S. Austr. 61 (1937) 248; Fl. S. Austr. pt 4 (1957) 
773, f. 1106. — Campsis pandorana Steen. Fl. 
Males. I, 4 (1948) xxi. — Fig. 37a-b. 

Malesian specimens: Often large liana, 20-30 m. 
Leaves 2-4(-6)-jugate; leaflets usually ovate- 
elliptic to oblong, mostly entire and acuminate, 
3-10 by 1 ',2-6 cm, glabrous, underneath with 
few or many scattered large crateriform glands, 
nerves not prominent above; midrib sulcate; 
lateral petiolules 0-10 mm; petiole without glands 
at the base above. Thyrses terminal, lateral or 
from old wood, c. (l-)5-20 cm, glabrous, rarely 
puberulous. Pedicels c. l l 2 -l 1 U cm, rarely longer. 
Calyx cupular, stunted or short-lobed, thin, c. 
2-3 mm. Corolla l-2(-3)cm long including the 
lobes, mostly rather inflated-tubular, the tube 
mostly ± twice as long as the lobes, glabrous 
outside, lobes mostly densely papillose-puberulous, 
mouth and tube inside bearded on the ventral side, 
light yellow, the zygomorphous limb and tube 
inside streaked or mottled red or purple dotted, 
without a hair-ring near the base of the stamens 
and no proper basal tube. Stamens included, at 
their base glandular-papillose, inserted very near 
the base of the tube. Ovary glandular-papillose. 
Capsule acute, c. (5-)9-12 by (l 3 /^-)2V2-3 by 
2-2Vjcm; valves coriaceous; dissepiment rather 
thick, 6V2-8 by l 3 / 4 -2 cm, the seed scars marginal. 
Seeds c. 2 l 2 -3 by 1 "2-2 cm including the hyaline 
wings. 

Distr. Central, N. & E. Australia, Tasmania, 
Lord Howe I., New Caledonia, N. Solomons 
(Bougainville), and East Malesia: New Guinea 
(incl. New Britain), Moluccas (Morotai, Halma- 
heira, Ambon, Ceram, Key Is.), and Lesser Sunda 
Is. (Lombok, Flores, Timor); 65 collections. 

Though Tecoma filicifolium, a juvenile form, was 
said to have come from Fiji, I have shown (1967, 
I.e.) that this hailed from New Caledonia. 

Sims claimed that Loddiges nurseries had 
received Bignonia pandorae from Norfolk I., 
but this seems to rest either on an erroneous localis- 
ation or on a cultivated source. 

Ecol. In Malesia in primary and secondary rain- 
forest, from sea-level up to c. 2000(-2400) m, 
getting distinctly scarcer upwards of 1350 m. Fl. 
Jan.-Dec.,/r. Oct. -Dec. 

Tax on. A quite well recognizable species in 
spite of a fair degree of variability. This is in part 
ontogenetic, the juvenile form having narrow, 
many-jugate, crenate, small leaflets; these are 
sometimes still found on odd twigs of mature- 
foliaged plants. 

The main variation is in Australia in the leaves, 
the rain-forest (type variety) form having ovate to 
elliptic 2-jugate leaflets, whereas in drier places 
2 4-jugate leaves occur with lanceolate leaflets 
(described as T. mconanthu), while in still more 
arid places the 2 6-jugatc leaves have almost linear 
leaflets (described as T. oxleyi and T. doratoxylon). 
The latter form may at times be scrambling, bushy 
or even erect (spcarwood bush) and carry racemose 
inflorescences. Though the typical representatives 



of these three forms are distinct, they are con- 
nected by a clear series of specimens with inter- 
mediary characters, which already induced Bailey 
to say that he named the three forms as varieties 
merely for convenience. Whether these forms are 
genetically different taxa (races) or merely pheno- 
typic forms can only be established by experiments. 
Bailey added that flowers of the type variety 
would emit a strongly disagreeable odour while 
T. meonantha would have fragrant flowers. This 
matter must be solved by field botanists. 

In Malesia the 3-4-jugate leaves are often 
narrower, sessile, and also more toothed than the 
2-jugate ones; it looks like a matter of lingering 
neoteny. 

At higher altitudes, 1300-2000 m, leaflets tend 
to be more coriaceous. In Schmutz 3178 the veins 
are by exception prominent above. 

The New Caledonian form seems to be a local 
race with small flowers and small roundish, dentate 
leaves. 

In rain-forest the leaflets are usually entire, or 
with a few coarse teeth towards the apex, and 
usually they have underneath a fair number of 
crater-like larger glands, in Australian specimens 
these are scarce or absent. 

Lateral petiolules are usually short (2-5 mm), 
but in Hyland 5092 they measure 17a cm. 

la. ssp. timorensis Steen. nov. ssp. 

Differt a speciminibus malayanis floribus com- 
parate magnis (2-3 cm longis, lobis incl.), stamini- 
bus atque stylo exsertis, ceterum ore atque tubo 
floris bar bis longis destitutis. — Typus: C. W. Kooy 
363 (L), pr. Temef, S. Central Timor, fl. 18-vii- 
1966, c. 800 m. 

Leaflets 2-4-jugate, without crateriform glands 
beneath. Calyx 2-4 mm, stunted, minutely 
5-mucronulate. Corolla 2-3 cm long incl. lobes. 
Mouth and tube not bearded inside, the puberulous 
papillae from the lobes extending in the tube in a 
lax way. Stamens and style exserted. 

Distr. Malesia: Lesser Sunda Is. (Timor); 4 
collections. 




Fig. 38. Range of the genus Vumlorea Spach. 

Figures above the hyphen indicate endemic species, 

those below the hyphen non-endemic species. 



178 



Flora Malesiana 



[ser. I, vol. 8 2 



Ecol. On limestone (once) and along ravine in 
mountain Eucalypt forest, c. 700-1000 m. Fl. 
March, May, July. 

Vern. Tufe, Dawan lang., non amisu, Niki-Niki, 
nonfuleh, Mt Mutis. 

Notes. Though undoubtedly P. pandorana, the 
Timor race deviates within the species by lacking 
the usual beard in the mouth and tube of the 
corolla and in the genus by exserted stamens and 
style. It is remarkable that the specimens from 
Lombok and Flores do not belong to this sub- 
species but agree with the Moluccan specimens. 

2. Pandorea stenantha Diels, Bot. Jahrb. 57 (1922) 
498; Steen. Nova Guinea 14 (1927) 302; Thesis 
(1927) 850; Bull. Jard. Bot. Btzg III, 10 (1928) 
197. — Fig. 39e-f. 

Large liana (stem to 2 l / 2 cm 0). Leaves (1— )2— 3- 
jugate, the leaflets coriaceous, ovate-oblong, acute, 
towards apex toothed, venation raised on both 
sides, above fine-tessellate, 6-8V2 by 3V2-4cm; 
rachis narrowly winged ; petiolules 0-8 mm ; 
petiole near the base above with one or few large, 
sometimes raised glands, sometimes also one at the 
articulation of the rachis. Thyrses axillary and 




Fig. 39. Pandorea montana (Diels) Steen. a. Leaf, 
b. leaflet, x l /a» c. reticulate venation above, d. 
inflorescence, x l j 2 . — P. stenantha Diels. e. 
Inflorescence, x l j 2 , f. tessellate venation of leaflet 
above (a, c NGF 13905, b Ledermann 9916, d 
NGF 21251, e Soegeng 360, f Van Royen & 
Sleumer 5816). 



terminal or on old wood, 10-15 cm, fine- 
puberulous. Pedicels 4-10 mm. Calyx 3-5 mm, 
campanulate, with 5 short, broad-deltoid lobes. 
Corolla tube yellow, the lobes white (ex typ.), 
tube c. 20 mm long and c. 5 mm wide, outside 
puberulous-papillose, the mouth and upper part 
of the tube bearded, lobes c. 3-4 mm; no hair-ring 
at the insertion of the stamens. Stamens included. 
Disk cupular. Ovary flattened, obconical-oblong, 
glandless. 

D i s t r. Malesia: New Guinea (Sepik Distr. : April 
R. ; Mt Cyclops ; SE. Irian : Ingembit) ; 3 collections. 

Ecol. Rain-forest, 125-800 m. Fl. June, Nov. 
Flowers once noted to be fragrant. 

Notes. Apparently a rare species; available 
flower material unfortunately rather inadequate. 
Van Royen noted : flowers white at base of tube 
pale purple; Soegeng: tube dirty yellow, lobes 
lilac. The petiolar glands were not mentioned by 
Diels in his brief description, but I noted them on 
the type (1927). 

3. Pandorea montana (Diels) Steen. nov. comb. — 
Tecomanthe montana Diels, Bot. Jahrb. 57 (1922) 
497; Steen. Nova Guinea 14 (1927) 296; Thesis 

(1927) 875, f. 3c; Bull. Jard. Bot. Btzg III, 10 

(1928) 204. — Fig. 39a-d. 

Slender liana, to finger-thick. Leaves 3-4(-5)- 
jugate, leaflets chartaceous to coriaceous, elliptic- 
oblong, acute, towards apex toothed or shallowly 
crenate, venation raised on both sides, but not 
fine-tessellate above, c. 2-5 1 / 2 by 1-2V 2 cm, rachis 
narrowly winged; petiole without basal glands; 
petiolules almost absent. Thyrses axillary and 
terminal, lax, almost glabrous. Pedicels filiform, 
1-2 cm. Calyx c. 7 mm, with 5 short, broad- 
triangular teeth. Corolla ± infundibuliform, 
20-25 mm long, the tube narrowed to the base, 
straight, limb zygomorphous, c. 8-10 mm wide at 
the mouth, the lobes 2-4 mm, bearded in the 
mouth and upper part of the tube, the lobes white 
to pale red or streaked red, the tube light brown 
or yellow flushed red outside, yellow inside, with a 
distinct stuppose hair-ring near the insertion of the 
filaments. Stamens included. Disk cupular. Ovary 
± conical, eglandular. Capsule flattened, 4-8V2 
by IV2-3 by V2-I cm, sessile to stiped and short- 
beaked; valves hard, almost coriaceous to woody, 
narrow boat-shaped ; dissepiment thick-coriaceous, 
3-10 by VU-l'^cm. Seeds l l l 4 -2 3 U by 1-2 cm. 

Distr. Malesia: East New Guinea (Sepik: 
Hunstein Mts: Lordberg, Hunsteinspitze, 1000- 
1350m, 4 coll.), loc. class.; Morobe Distr.: Mt 
Kaindi, 7°25', 146°45' E, 12 coll.; Mt Giluwe, 
6°10', 144° E. 

Ecol. Mountain rain-forest, often mossy, 
forest edges and regrowths, 1000-1350, 2100- 
2450 m. Fl. Febr., May-June, Aug.-Nov., fr. 
Aug.-Sept., Dec. 

Notes. The thyrsoid inflorescence and flower- 
size point to congenerity with Pandorea; luckily I 
had made in 1927 a small drawing of the type and 
observed several flower details on the original 
material not mentioned by Diels in his brief 
description. Unfortunately almost all duplicates at 
L lack flowers, by careless distribution. The flower 
colour noted on labels varies rather considerably : 
tube cream, yellow, golden brown or red outside, 
lobes white, cream, pale purple, streaked red. 



1977] Bignoniaceae (van Steenis) 179 

Doubtful & Excluded 

In this list are combined all names of Malesian and SE. Asian plant names which are excluded or of 
which the identity is uncertain or of which I have not seen the types. 

Bignonia albida Bl. Verh. Bat. Gen. 9 (1823) 195; Steud. Nomencl. 2 (1841) 204 = Aeschynanthus 
albidus (Bl.) Steud. (Gesneriaceae). 

Bignonia angustifolia Bl. Verh. Bat. Gen. 9 (1823) 194; Cat. Hort. Bog. (1823) 82 = Aeschynanthus 
angustifolius (Bl.) Steud. {Gesneriaceae). 

Bignonia comosa Roxb. [Hort. Beng. (1814) 95, nom. semi-nudum] Fl. Ind. ed. Carey 3 (1832) 103 ; DC. 
Prod. 9 (1845) 144; Miq. Fl. Ind. Bat. 2 (1858) 751. — Spathodea comosa G. Don, Gen. Syst. 4 (1838) 
222, said to come from the Moluccas, is according to the type (Herb. Martius, in BR) Clerodendron 
lanuginosum Bl. 1825 (Verbenaceae). The detached fruit on the sheet was not described and belongs to 
some SE. Asian Bignoniaceae. 

Bignonia compressa Lamk, Encycl. 1 (1785) 424; G. Don, Gen. Syst. 4 (1838) 220, said to come from 
the East Indies, is according to Perrier de la Bathie from Madagascar (Fl. Madag. 178, 1938, 59) 
(Colea decora DC. Prod. 9, 1845, 241) = Rhodocolea racemosa H. Perrier. 

Bignonia fraxinoides Perrottet, Mem. Soc. Linn. Paris 3 (1824) 102, nom. semi-nudum, said to grow in 
East Java, is cf. Steen. Blumea 15 (1967) 146 probably not from Java,; the name ought to be discarded 
entirely. 

Bignonia glauca Wall. Cat. 6506, nomen, non Decne 1844. The type at Kew, a sterile specimen, was 
annotated "perhaps not Bignoniaceae" by C. B. Clarke. It has recently further been annotated "prob. 
Meliaceae" by Alan Radcliffe-Smith. 

Bignonia hirsuta Lamk, Encycl. 1 (1785) 422; Willd. Sp. PI. 3 (1802) 299; G. Don, Gen. Syst. 4(1838) 
225. — Tecoma hirsuta DC. Prod. 9 (1845) 173, 222. 

Said to have come from "lTndes" ; leaves opposite, digitate, with 5 leaflets, stalked and with petiolules ; 
leaflets oblong, cuneate at base, emarginate at apex, downy, slightly pubescent beneath. Flowers small, 
curved, short reddish-yellow hairy. Calyx truncate, with 4 minute teeth. Stamens 4, exserted. 

Vitex (Verb.) might be involved, but all its Indo-Malesian species have acuminate leaves and in Vitex 
the corolla is not curved and if hairy the corolla is greyish. It might possibly be an American plant, 
Tabebuia, or allied to that. 

Lamarck described the specimen from herb. Jussieu. Dr. H. Heine made elaborate but unfortunately 
unsuccessful attempts in herb. Jussieu and Lamarck to locate the specimen in the Paris Herbarium; there 
is also no trace of it at Geneva ; De Candolle did not see any material. 

Bignonia laeta Wall. Cat. 6505 A, cf. Clarke, Fl. Br. Ind. 4 (1884) 376, is according to Sprague, 
Kew Bull. (1919) 306 = Dolichandrone serrulata Seem. 

Bignonia longiftora Reinw. msc. ex De Vriese, PI. Ind. Bat. Or. (1856) 9, nomen = Aeschynanthus 
longiflorus (Bl.) DC. {Gesneriaceae). 

Bignonia macrostachya Wall. [Cat. 6504, nomen) ex G. Don, Gen. Syst. 4 (1838) 221 ; DC. Prod. 9 
(1845) 166. Mr. R. K. Brummitt, Kew, kindly remarked on this (in litt. 15-vm-1975) that "Wallich 
8504 consists of two rather long inflorescences in bud only (one corolla almost open) and a fairly stout 
piece of stem, and bears no leaves, open flowers or fruits. It has been annotated ''Bignonia macrostachya 
Wall, (and of G. Don & DC.)' by C. B. Clarke, but has no more recent identification. It seems to me to 
be fairly clearly referable to Pajanelia longifolia (Willd.) K.Sch. My opinion seems to be supported by a 
specimen laid away in the main herbarium under this species in a red folder (though it is not obvious what 
it is supposed to be a type of), collected in Khasiya by Griffith and labelled "Bignonia macrostachya 
Wall. Cat. 6504 & - rostrata Wall. Herb. 6503A'. The specimen Wallich 6503A does indeed also 
seem to be this species. The citation of Wallich 6505 by G. Don in validating the name B. macrostachya 
is presumably an error for 6504. 

Bignonia moluccana DC. Prod. 9 (1845) 144; Miq. Fl. Ind. Bat. 2 (1858) 751. — B. discolor A. Rich. 
Sert. Astrol. (1834) xxix, non R.Br. 1814, said to come from the Moluccas. The description would tally 
with Gmelina asiatica Lour. ( Verbenaceae), but a sheet with an original label (in P) was identified by 
E. Bureau as Bignonia capreolata L. which does not agree with the description. Confusion with labels and 
specimens must have taken place and the name should be discarded. Cf. Steen. Blumea 15 (1967) 146. 

Bignonia purpurea Thunb. Fl. Jav. (1825) 15, nomen. Of unknown identity, not mentioned by Juel. 

Bignonia ramiflora Decne, Nouv. Ann. Mus. Paris 3 (1834) 381, repr. Herb. Timor. Descr. (1835) 53. — 
? Bignonia colei G. Don, Gen. Syst. 4 (1838) 221. — Colea ramiflora DC. Prod. 9 (1845) 241 ; Miq. Fl. Ind. 
Bat. 2 (1858) 759. — Colea colei M. L. Green, Stand. Sp. Nom. Cons. (1926) 55-63; Steen. Bull. Jard. 
Bot. Btzg III, 10 (1928) 277, excl. syn. alter. — Colea timorensis in sched., in syn., ex Perrii r. 

This rests on a mislocalized specimen of Poivre in herb. Jussieu from Madagascar, and is according to 
Perrier de la Baihie, Ann. Mus. Col. Marseille 46 (1938) 43 = Rhodocolea racemosa H. Perrier. 

Bignonia ternatea Rusw. ex de Vriese, Reinwardt's Reize (1858) 495, 644, nomen = Dichrotrichum 
ternateum Reinw. ex Di Vriim 

Bignoniacea incerta: Zoll. Syst. Verz. Heft 3 (1855) 53-54, based on Zollinger 2214 = Wightia 
borneensis Hook./, ssp. otlolandcri (Koord.) Steen. (Scrophulariaceae). 

Colea aherrans Baii.i . Hull. Soc. Linn. Paris I (1889) 687 rests on a specimen said to have been collected 
by Poivre in Timor, but came from Madagascar according to Pirriir de la Haiku, Ann. Mus. Col. 
Marseille 46 (1938) 28, and = Rhodocolea racemosa Perrier var. humblotiana \\. Perrier. 

DoIk handrone falcata (Wai i . r> l)( ) Sum.: F.-Vii.l. Nov. App. (1880) 151. According to Mikrii i , 
En. Philip. 3 (1923) 445, obviously an erroneous record from the Philippines of this Asian species. 



180 Flora Malesiana [ser. I, vol. 8 2 

Dolichandrone tulipifera Bth. in B. & H. Gen. PI. 2 (1876) 1046; F.-Vill. Nov. App. (1880) 151 ; Merr. 
En. Philip. 3 (1923) 445. This is an erroneous, non-existing combination for Spathodea tulipifera G. Don 
which was used by F.-Villar for the tulip tree, Spathodea campanulata, which he saw cultivated at Manila. 

Hadongia eberhardtii Gagn. Not. Syst. 14 (1950) 30, from Indo-China, is according to Vidal, Bull. Soc. 
Bot. Fr. 106 (1959) 352 a cultivated specimen of Citharexylum spinosum L. (Verbenaceae). 

Hausmannia mollis K.Sch. ex Steen. in sched.; Thesis (1927) 902; Hausemannia mollis F.v.M. sphalm. 
Ind. Kew. Suppl. 1 (1906) 16 (Hausemannia mollis K.Sch.) = Archidendron molle (K.Sch.) De Wit 
(Leguminosae). 

Markhamia cauda-felina (Hance) Craib: Sprague, Kew Bull. (1919) 310. — Dolichandrone sp. Ceron, 
Cat. PI. Herb. Fl. For. Filip. (1892) 127. Now considered to be Markhamia stipulata (Wall.) Seem. var. 
cauda-felina (Hance) Santisuk, cf. Thai For. Bull. Bot. 8 (1974) 15. 

The collection on which this was based is Vidal 3398 (K), from Montufar, Albay Prov., Luzon. Merrill 
noted (J. Am. Arb. 35, 1954, 154) that it was possibly occasionally introduced for forestry purposes by 
Vidal from S. China. As no later collections were ever made its cultivation seems to have been ephemeral. 

Stereospermum cylindricum Pierre ex P. Dop, Fl. Gen. I.-C. 4 (1930) 581 , a species from Indo-China and 
Thailand, was mentioned by Dop, I.e. 582 to occur in Malaya, but on what evidence is unclear. I found no 
sheets in Paris to corroborate this. 

Tripinna tripinnata Lour. Fl. Coch. (1790) 391 . — Tripinnaria cochinchinensis Pers. Syn. 2 (1807) 173. — 
Tripinnaria asiatica Spreng. Syst. 2 (1825) 842, taken for a Bignoniacea by several authors. According to 
Merrill, Comm. Lour. (1935) = Vitex tripinnata (Lour.) Merr. (Verbenaceae). 

Cultivated Bignoniaceae 

There are quite a number of Bignoniaceae cultivated in Malesia ; they stem from all parts of the 
tropics. Frequently they hardly set any seed. Tecoma stans does so profusely and this has led to 
its naturalization. Jacarandas also set seed but did not naturalize. 

As Bignoniaceae are often very showy plants and are largely tropical there is no end to their 
introduction. Therefore the survey given below may be or at least become incomplete. 

It should also be remembered that cultivated plants are often neglected by botanical explorers 
and are mostly scantily represent in herbaria. 

These introduced species have mostly not been critically studied by me, but it is assumed that 
their names are correct. I acknowledge with great thanks the loyal collaboration of the late Mr. 
N. J. Sandwith (Kew) who formerly named at my request certain introduced species, and of 
Dr. A. H. Gentry (Missouri Botanical Garden, St. Louis) who was so kind as to check this 
appendix. 

Some papers or appendices are dedicated solely to cultivated Bignoniaceae or have taken them 
up and often give keys and illustrations: 

Backer, C. A. & Bakhuizen van den Brink Jr, R. C. 1965. Flora of Java 2: 534-542. 
Chatterjee, D. 1948. A review of Bignoniaceae of India and Burma. Bull. Bot. Soc. Beng. 2: 

75-79. 
Fabris, H. A. 1959. Las plantas cultivadas de la Republica Argentina. Bignoniacas. Inst. Bot. 

Agr. 10, fasc. 173: 57 pp., 25 fig. 
Gentry, A. H. 1973. Ann. Mo. Bot. Gard. 60: Flora of Panama, part IX, fam. 172: 781-977, 41 

fig. 
Heine, H. 1976. Flore de Nouvelle-Caledonie 7: 91-93. 

Herklots, G. 1976. Flowering tropical climbers: 63-74, fig. 80-101, col. pi. 5-7. 
Holttum, R. E. 1941. The Bignonia family in Malayan gardens. M.A.H.A. Mag. 11: 3-11. 
Santisuk, T. 1974. Bignoniaceae. Thai For. Bull. Bot. 8: 1-46. 
Sykes, W. R. 1966. Studies of cultivated plants in New Zealand. 1. Bignoniaceae. New Zeal. 

D.S.I.R. Inf. ser. 54: 63 pp., 25 fig. 

artificial key to cultivated bignoniaceae in malesia 

1 . Climbing plants. 
2. Leaves at least 2-jugate. Tendrils absent. 

3. Stamens exserted. Corolla narrow-tubular, scarlet or sulphur-yellow Tecoma capensis 

3. Stamens included. Corolla not narrow-tubular, lavendar or white streaked with carmine. 
4. Calyx truncate, at most 6 mm, including minute teeth, not inflated. 
5. Corolla c. 4-5 cm long, white with crimson throat, the tube puberulous outside. Calyx c. 6 mm. 

Pandorea jasminoides 



1977] Bignoniaceae (van Steenis) 181 

5. Corolla ( 1 — )2— 3 cm long, pale yellow, lobes purple dotted or streaked, the tube glabrous outside. 

Calyx 2-3 mm Pandorea pandorana 

4. Calyx campanulate, inflated, white, distinctly 5-lobed, P/2 cm long . . . Podranea ricasoliana 
2. Leaves 1-jugate. Tendrils in a number of leaves present. 
6. Corolla lobes valvate; tube narrow, without a distinct basal tube. Stamens exserted. Flowers 

bright orange Pyrostegia venusta 

6. Corolla lobes imbricate. 
7. Calyx spathaceous, thin, 3 cm long. Corolla very large, rose-purple, (5-)7-9 cm. Disk absent. 

Phryganocydia corymbosa 
7. Calyx regular, not spathaceous. Disk present. 
8. Inflorescence, calyx and outside of corolla tube densely hairy. 

9. Calyx 5-6 mm long, 6-8 mm wide at the mouth. Inflorescence pauciflorous, almost a raceme, 
with 2-4 pairs of opposite flowers. Tendrils branched Pithecoctenium cynanchoides 

9. Calyx 3-4 mm long, 2" 2 -4 mm wide at the mouth. Thyrse rich-flowered, the flowers in triad 
cymes. Tendrils unbranched Arrabidaea mollissima 

8. Inflorescence, calyx and outside of corolla tube glabrous. 

10. Tendrils undivided. Leaves obovate with long-cuneate base, the two basal nerves straight, 
running up over halfway the blade, in the narrow angle with the midrib a large dark coloured 
gland field. Pseudostipules present Saritaea magnifica 

10. Tendrils 3-parted. Leaves not obovate, and no such gland field in the narrow angle at the base. 
11. Corolla yellow, outside of lobes glabrous or lepidote. Plant without garlic odor. 
12. Tendril with 3 claws. Pseudostipules scale-like. Calyx thin, broadly campanulate 

Macfadyena unguis-cati 
12. Tendril 3-fid. Pseudostipules foliaceous, 5— 7(— 1 5) mm in diameter. Calyx coriaceous, cupular. 

Anemopaegma chamberlaynii 
11. Corolla pale mauve to pale purple, outside of lobes puberulous. Plant smelling of garlic. 

Pachyptera hymenaea 
1 . Erect shrubs or trees. 

13. Leaves in scattered fascicles. Fruit indehiscent, hard-shelled with fleshy pulp. 
14. Leaves simple Crescentia cujete 

14. Leaves 3-foliate, petiole winged Crescentia alata 

13. Leaves decussate or in whorls. Fruit dehiscent or fleshy and without a hard shell. 

15. Leaves simple. 

16. Leaves elliptic on a long, slender petiole. Thyrses terminal. Capsule linear with long-hairy, linear 
seeds. Unarmed tree Catalpa longissima 

16. Leaves small, obovate, tapering into the base; no proper petiole. Flowers ramiflorous. Fruit an 
oblong berry. Twig nodes with a pair of thorns Parmentiera aculeata 

15. Leaves compound. 

17. Leaflets 3, sessile, articulated on top of a winged petiole. Fruit fleshy. 

18. Branchlets usually with 2 short, ascending thorns at most nodes. Fruit costate and curved, less 
than 1 7 cm long Parmentiera aculeata 

18. Branchlets unarmed. Fruit neither curved nor costate, usually more than 30 cm long, candle-like. 

Parmentiera cereifera 
17. Leaflets 5 or more. 

19. Leaves digitately compound. Leaflets 5. 

20. Leaflets stellate-hairy beneath. Flowers yellow. Calyx rusty stellate-tomentose 

Tabebuia chrysantha 
20. Leaflets lepidote, otherwise glabrous. Flowers pink. Calyx lepidote. 
21. Leaflets acuminate or sharply acute. Inflorescence usually many-flowered. Mature capsule 

more than 22 cm long Tabebuia rosea 

21. Leaflets obtuse. Inflorescence few-flowered. Mature capsule usually less than 15 cm long 

Tabebuia pallida 
19. Leaves pinnate. 
22. Leaves 2-pinnate. 

23. Corolla infundibuliform, lilac. Capsule broad-ellipsoid, with woody valves. Staminode longer 
than the stamens, glandular-pubescent, especially at apex. 
24. Corolla glabrous, distinctly sigmoid. Leaflets acute, with oblique base. 

Jacaranda obtusifolia ssp. rhombifolia 
24. Corolla densely short-hairy, not sigmoid. Leaflets cuspidate, base hardly oblique 

Jacaranda mimosifolia 
23. Corolla salvcr-shapcd, white, the tube 6 8 cm long, 2 mm wide. Capsule linear. Staminode 

absent Millin^tonia hortensis 

22. Leaves 1 -pinnate. 
25. < ,il.\ CMrae, 2 7cm long. Corolla wide-campanulatc, coarse and large. 
26. Calyx irregularly lobed, 2 3 cm. Flowers inside dark red, nocturnal, in long, pendent racemes. 

Kerry massive, sausage-shaped Kinelia africana 

26. ( alyx spathaceous, 4 7 cm. I lowers orange-red, in erect terminal thvises, diurnal. Capsule 

dehiscent, with winged seeds Spathodca campanulata 

25. Calyx short 5-lobcd, 5-6 mm. Flowers not coarse. 



182 



Flora Malesiana 



[ser. I, vol. 8 2 



27. Corolla infundibuliform, yellow, with included stamens. Leaflets lanceolate, serrate, some- 
times deeply incised Tecoma stans 

27. Corolla narrow-tubular, usually orange red, sometimes sulphur-yellow, with exserted stamens. 
Leaflets ovate, dentate Tecoma capensis 



Anemopaegma chamberlaynii (Sims) Bur. & K.Sch. 
Fl. Bras. 8, 2 (1896) 128; Herklots, Fl. Trop. 
Climb. (1976) 65, f. 82. — Bignonia chamberlaynii 
Sims, Bot. Mag. (1820) t. 2148. — Bignonia 
scandens Vell. Fl. Flum. 6 (1825) 232, t. 22. — A. 
scandens Mello ex K.Sch. in E. & P. Nat. Pfl. 
Fam. 4, 3b (1894) 215; Back. & Bakh./. Fl. Java 2 
(1965) 536. 

Glabrous. Pseudostipules foliaceous, ovate to ± 
orbicular, 5-7(-15)mm. Leaflets ovate-oblong, to 
lanceolate-oblong, acute, 5-14 by 2V2-5V2 cm. 
Tendrils 3-fid. Flowers in 2-8-flowered axillary 
racemes. Calyx campanulate, truncate, 7-8 mm. 
Corolla 4-5 cm, pale yellow. 

Distr. Brazil, introduced in East Java as an 
ornamental, at Malang and Kali Baru (Besuki), 
250-600 m; also seen from Rangoon, Burma 
(Dickason 6660). All Asian material has smallish 
ovate leaflets, 5-6 by 2 l l 2 -VI 2 cm and smallish 
acute pseudostipules 5-7 mm long; in America 
both can obtain much larger size. 

Arrabidaea mollissima (H.B.K.) Bur. & K.Sch. 
Fl. Bras. 8, 2 (1896) 46; Seibert, Carnegie Ins., 
Wash. 522 (1940) 406; Dugand, Caldasia 3 (1945) 
255. 

Leaves patent, lax hairy, the longer hairs mostly 
gland-tipped; leaflets ovate, acuminate, 4-12V 2 by 
2V2-7V2 cm. Flowers pink to rose, 3V 2 -5 cm, 
mouth whitish with yellow spot. 

Distr. Mexico and Central America to Colum- 
bia and Venezuela; cultivated in the vicinity of 
Manila. 

Catalpa longissima (Jacq.) Dum. Cours. Bot. Cult. 
2 (1802) 190; Sims, Bot. Mag. (1808) t. 1094; 
Sandwith, Rec. Trav. Bot. Neerl. 34 (1937) 228; 
Adams, Fl. PI. Jamaica (1972) 669; Little Jr., 
Trees Puerto Rico & Virgin Is. 2 (1974) 890, f. 700. 

— Bignonia longissima Jacq. En. PI. Carib. (1760) 
25. — Macrocatalpa longissima Britton, J. N.Y. 
Bot. Gard. 19 (1918) 8. 

Tree up to 30 m, occasionally deciduous. Petioles 
slender; blades ovate-lanceolate, acute, (3-)5-l 1 by 
(lV2-)2-4cm. Flowers in small panicles, white, 
pinkish on the lobes, yellow in mouth with purple 
markings, 2*/2-3 cm long. Calyx 2-cleft. Capsule 
35-75 cm by 4 mm. 

Distr. Jamaica, Hispaniola, Martinique, intro- 
duced for ornamental or forestry purposes in other 
tropics, e.g. in the Marianas (Guam, Saipan) and 
the Philippines (Luzon: Lamao For. Res.). 

Vern. French oak, Haitian oak, Mast-wood, 
Yoke-wood, E. 

Crescentia alata H.B.K. Nov. Gen. Sp. 3 (1819) 
158; F.-Vill. Nov. App. (1880) 151; Vidal, 
Sinopsis Atlas (1883) 35, t. 73, f. C; Merr. Fl. 
Manila (1912) 430; Sp. Blanc. (1918) 350; En. 
Philip. 3 (1923) 447 ; Standley, Trees Shrubs Mex. 
(1926) 1324; Back. & Bakh./. Fl. Java 2 (1965) 
542; Gentry, Ann. Mo. Bot. Gard. 60 (1973) 829. 

— C. trifolia Blanco, Fl. Filip. (1837) 489; DC. 



Prod. 9 (1845) 247; Blanco, Fl. Filip. ed. 3. 2 
(1878) 271, t. 327. — Otophora paradoxa Bl. 
Rumphia 3 (1847) 146; Mio. Fl. Ind. Bat. 1, 2 
(1859) 560 (L). — Parmentiera alata Miers, 
Trans. Linn. Soc. Bot. 26 (1870) 166. 

A crooked tree, 5-14 m, to 25 cm 0. Leaves in 
scattered fascicles on the twigs, with a winged 
petiole, 2V2-Hcm; leaflets brittle, lanceolate- 
obspathulate, sessile, l-4V2cm. Flowers 1-2 
rami- and cauliflorous; calyx 2-lobed to the base; 
corolla brownish with brown-purple venation, 
rank-scented, with a transversal fold, 4-6 cm. 
Fruit with a hard shell, ± globular, 5-10 cm 0. 

Distr. Mexico to Costa Rica, from Mexico 
early introduced by the Spaniards via Guam in the 
Philippines, also in Java (rare) and Rabaul (New 
Britain). 

Vern. Philippines: cruz-cruzan, Tag., hoja cruz, 
Spanish. 

The pulp is in Mexico sometimes used as a 
medicine ; the shells are less in demand than those 
of Crescentia cujete, but used for the same pur- 
poses. No fruits seen from Malesia. 

Crescentia cujete Linne, Sp. PI. (1753) 626; 
Standley, Trees Shrubs Mex. (1926) 1324; Steen. 
Thesis (1927) 1010; Bull. Jard. Bot. Btzg III, 10 
(1928) 274; Seibert, Carnegie Inst. Wash. 522 
(1940) 383; Back. & Bakh. /. Fl. Java 2 (1965) 
542; Gentry, Ann. Mo. Bot. Gard. 60 (1973) 831. 
— C. ovata Burm. /. Fl. Ind. (1768) 132, nom. 
semi-nudum (G). 

Crooked tree to 10 m, 30 cm 0. Leaves in scat- 
tered bundles on the rough twigs, obspathulate, 
sometimes short-acuminate, without petiole, up to 
26 by 7 x / 2 cm. Flowers solitary or in pairs on the 
twigs, of a musty odor ; calyx 2-lobed to the base ; 
corolla 4-7 cm, dirty white or pale greenish, 
purplish veined, finally turning dull purplish, the 
wide tube with a transversal fold. Fruit broad- 
ellipsoid to globular, 1 3-20 by up to 30 cm, indehis- 
cent, with a hard shell. 

Distr. Central America, very widely and early 
distributed in the American and other tropics, 
throughout Malesia, in the lowland, grown in 
lawns, parks and used for hedges. 

Vern. Tabu kaju, S. Sum., bila, Djakarta, 
sekopal, sikadel, J, bila radja, Kangean, bila 
bilanda, Makas., calabassa, Moluccas, ber(e)nuk, 
S, bua no, Ternate, buwano, Halmaheira, Calabash 
tree, E. 

The pulp is sometimes used for medicinal 
purpose and the hard shells are commonly in use 
for drinking cups, vessels, and carving. 

The flowers are bat-pollinated. Cf. Porsch, Oest. 
Bot. Z. 80 (1931) 31^14, t. 9-10. 

I saw the type of C. ovata Burm. /. which was 
described from Java and which was in Index 
Kewensis reduced to C. cucurbitina L. (now 
Amphitecna latifolia (Mill.) Gentry); the type 
consists of 3 leaves and a single damaged flower; 
the sheet carries no name, but the note 'Kalbas, 
4 stam., 1 pistile'. 



1977] 



Bignoniaceae (van Steenis) 



183 



Jacaranda mimosifolia D. Don, Bot. Reg. 8 (1822) 
t. 631. — y. ovalifolia R.Br. Bot. Mag. 49 (1822) 
t. 2337. — J. acutifolia (non H. & B.) auct. ; Steen. 
Bull. Jard. Bot. Btzg III, 10 (1928) 270; Back. & 
Bakh./ Fl. Java 2 (1965) 239. 

Distr. Tropical America, Bolivia to NW. 
Argentina. Seen from W. Java, Hawaii, S. Africa, 
Madagascar, Congo. Perhaps not distinct from the 
Peruvian J. acutifolia H.B.K. 

Suitable as a roadside tree and in parks. Flowers 
often when leaves are shed. 

The names of Don and Brown were published 
on the same day on material from the same source; 
a unique nomenclatural case! 

Jacaranda obtusifolia H.B.K. ssp. rhombifolia 

(G. F. W. Meijer) Gentry, Mem. N.Y. Bot. 
Gard., in the press. — J. rhombifolia G. F. W. 
Meijer, Fl. Esseq. (1818) 213. — J. filicifolia 
D. Don, Edinb. Phil. J. 9 (1823) 266; Steen. 
Bull. Jard. Bot. Btzg III, 10 (1928) 269; Holttum, 
M.A.H.A. Mag. 3 (1933) 188, fig. ; Back. & Bakh. 
/. Fl. Java 2 (1965) 239. 

Distr. Northern South America, Venezuela to 
Guiana. Seen from Malaya, Java (W. Java, also 
Semarang and Malang), and Borneo (Sandakan, 
Kuching). 

Good roadside tree, and for large gardens and 
parks. 

Kigelia africana (Lamk) Bth. in Hook. Niger Fl. 
(1849) 463; Sprague, Fl. Trop. Afr. 4, 2 (1906) 
536; Merr. Fl. Manila (1912) 430; En. Philip. 3 
(1923) 444; H. Heine in Hutch. & Dalz. Fl. W. 
Trop. Afr. ed. 2, 2(1963) 385; Merxm. & Schreib. 
Prod. Fl. SW. Afr. fam. 128 (1967) 3. — Bignonia 
africana Lamk, Encycl. 1 (1785) 424. — Crescentia 
pinnata Jacq. Coll. 3 (1789) 203, t. 18. — Tecoma 
africana G. Don, Gen. Syst. 4 (1838) 224. — K. 
pinnata DC. Prod. 9 (1845) 247; Koens, Trop. 
Natuur 1 (1912) 167, 6 f. — K. aethiopica (Fenzl) 
Decne in Deless. Ic. Sel. PI. 5 (1845) 39, t. 93; 
Srns. Bull. Jard. Bot. Btzg III, 10 (1928) 275; 
Back. & Bakh./. Fl. Java 2 (1965) 542. — See for 
further synonyms Heine, vide supra. 

Widely branched tree, to 20 m. Leaves decussate 
or in whorls of 3-4, up to 50 cm; leaflets oblong, 
entire or serrate distally, glabrous or hairy on 
nerves, to 20 by 6 cm. Flowers nocturnal, coarse, in 
terminal, pendent, narrow panicles up to 2 m long; 
calyx greenish, 2 3 cm, 2-lipped; corolla outside 
yellowish, veined, inside dark wine-red, 5-10 cm; 
basal tube as long as the calyx or longer. Berry 
sausage-like, 25 50 by 7 1 2 -l5cm, often on still 
flowering panicles 

Distr. Africa, widely cultivated in other tropics 
as an ornamental tree in parks and along roads, in 
Malesia not rare. 

Ilir.i /.< . concluded that the genus consists of 
one species only as the species distinguished by 
Stapi are racial and grading. 

The calyx contains much watery slime in bud 
[d KOOftD. Ann Jard, Hot. Ht/K 14, 1897, 407 
411). (he flowers are in Malesia invariably visited 
by bats. (f. FAIRCHILO, Imp. Gard, Hull. July 
I96K, 5, Harms Ac Hakim, The Nigerian Held 40 
151 158; J. West Afr Sc. Assoc. 4 (1958) 25 30 
Natuur I (1912) 167, I. I 6 
Mc(as-.,j bomb. Net. Hurt. Soc <mi)4fc7 47i, 



3 f. They are, however, also frequented by hawk- 
moths and Harris & Baker ll.ee. concluded that 
bats are not essential pollinating visitors. 

Macfadyena unguis-cati (L.) A. Gentry, Brittonia 
25 (1973) 236; Ann. Mo. Bot. Gard. 60 (1973) 874. 

— Bignonia unguis-cati Linne, Sp. PI. (1753) 623. 

— Doxantha unguis-cati Miers em. Rehder, Mitt. 
Deut. Dendr. Ges. (1913) 262; Herklots, Fl. 
Trop. Climb. (1976) 66, f. 87. — Bignonia tweediana 
Lindl. Bot. Reg. 26 (1840) t. 45, non Griseb.; 
Buysman, Flora 107 (1915) 361, cult, in Java. — ? 
M. dentata Bur. & K.Sch. Fl. Bras. 8, 2 (1897) 291 ; 
Steen. Bull. Jard. Bot. Btzg III, 10 (1928) 186; 
Back. & Bakh./. Fl. Java 2 (1965) 538. 

Leaves very variable, those of juvenile plant 
appressed to substratum, very small ; later to 5 by 
3'/2 cm, ovate, dentate, but in other forms elliptic- 
oblong and hardly dentate. Calyx irregularly 
lobed to spathaceous or subspathaceous. Corolla 
4'/ 2 -8 cm. 

Distr. Mexico to Brazil and N. Argentina, some- 
times cultivated in Malesia. 

Specific characters in this genus seem to be vague ; 
those used to distinguish M. uncata (Andr.) 
Sprague & Sandw. from M. unguis-cati by Gentry 
(Ann. I.e. 871) he declared himself (Brittonia 
I.e. 236) as inconstant. The one specimen I have 
seen from Java agrees with the plate of M. dentata 
but for the non-spathaceous calyx. Shape of calyx, 
margin of leaves and length of corolla seem to be 
very variable in this species which has alrez dy a very 
large synonymy (Gentry, Ann. I.e. 871). Does not 
fruit in Java; propagated by suckers and cuttings. 

Millingtonia hortensis L. / Suppl. (1781) 291. See 
for a full treatment p. 133. 

Distr. SE. Asia, probably also native in the 
Lesser Sunda Islands. 

Suitable as a tree for roadsides and parks. 

Pachyptera hymenaea (DC.) Gentry, Brittonia 25 
(1973) 236; Ann. Mo. Bot. Gard. 60 (1973) 888, 
with full synonymy. 

Glabrous liana. Vegetative parts smelling of 
garlic. Pseudostipules bract-like, l'^rnm. Leaflets 
triplinerved, ovate-oblong, short-acuminate, 372 _ 
7 by P/2-3 cm; venation prominent on both sides. 
Flowers in short axillary racemes. Calyx tube 
4-5 mm, with minute prominent pustular glands. 
Basal tube of corolla twice as long as calyx, in all 
3-4 cm. Anthers glabrous. Fruit flattened, 16-22 by 
IV4-I72 cm; valves with a central rib. Seeds 
1-1 V2 by 3-3'/2cm, incl. the membranous wing. 

Distr. A common liana ranging from Mexico to 
Brazil of tropical dry forest, sporadically also in 
moist forest, introduced in the Philippines (Manila, 
Mindanao) and E. Java (Surabaya). 

Another species may also he cultivated which is 
m flower not easy to distinguish from /'. hymciuica, 
viz P. ulliacea (Lamk) GENTRY : this latter species 
has a short oblong fruit with thick, corky wingless 

The genus Pseudocalytimti Sami- A; Ki m m. has 
been reduced to Pachyptera a.ik by Qbnttc 

I'andoreu jasminciidis (I indi ) K.S< 11. in I. Ac P. 
Nat. I'll. lam. 4, 3b (1894) 230; Shin I hesis 



184 



Flora Malesiana 



[ser. I, vol. 8 2 



(1927) 847; Bull. Jard. Bot. Btzg III, 10 (1928) 195; 
Proc. R. Soc. Queensl. 41 (1929) 48; Back. & 
Bakh. /. Fl. Java 2 (1965) 538; Herklots, Fl. 
Trop. Climb. (1976) 69, f. 90. — Tecoma jasmi- 
noides Lindl. Bot. Reg. (1939) t. 2002. 

Fairly tall climber; leaflets 4-7(-9), lanceolate, 
blunt, 2V 2 -5 by 1-2 cm. Corolla white streaked 
with carmine in the mouth, 4-5 cm long, short 
hairy. 

Distr. NE. Australia, not rarely cultivated in 
the tropics and also subtropics (Mediterranean, 
N. New Zealand), in Malesia rarely cultivated 
{e.g. West Java), in the temperate zone in green- 
houses. 

Propagated by cuttings ; seeds very rare. 

Pandorea pandorana (Andr.) Steen. Bull. Jard. 
Bot. Btzg III, 10 (1928) 198; Back. & Bakh./ 
Fl. Java 2 (1965) 538. For full references see p. 176. 

Distr. East Australia, New Caledonia, Lord 
Howe I., Solomons (Bougainville), New Guinea, 
Moluccas, and Lesser Sunda Islands. 

In Malesia an unfrequent ornamental, but culti- 
vated here and there through the tropics and sub- 
tropics (also in the Mediterranean and N. New 
Zealand). 

Propagated by tjankoks (marcotting) and cut- 
tings. 

Parmentiera aculeata (H.B.K.) Seem. Bot. Voy. 
Herald (1854) 183; Seibert, Carnegie Inst. Wash. 
522 (1940) 385; Gentry, Ann. Mo. Bot. Gard. 60 
(1973) 899. — Crescentia edulis Desv. J. Bot. 4 
(1814) 112. — Crescentia aculeata H.B.K. Nov. 
Gen. Sp. 3 (1819) 158. — P. edulis DC. Prod. 9 
(1845) 244 (heterotypic with Crescentia edulis 
Desv.). 

Tree 7-8 m. Leaves articulated with very hard, 
thorny extension of the nodal bark, the latter 
remaining a permanent, ascending, sharp thorn; 
leaflets elliptic, narrowed at both ends, 2-5 by 
1-3 cm; axillary fascicled leaves mostly simple. 
Flowers 1-several together, terminal, axillary or on 
branches or stem. Calyx spathaceous, 3-5 cm; 
corolla white, 6-7 cm. Berry pendent, cylindric, 
curved, costate with thick ribs, 8-17 cm by over 
3 cm 0. 

Distr. S. Mexico to northern Central America, 
cultivated elsewhere in the tropics; in Malesia rare: 
W. Java, Luzon, also seen from Cairns (N. 
Queensland). 

Young sterile offshoots have no thorns and 
possess coarsely dentate leaflets. 

Parmentiera cereifera Seem, in Hook. J. Bot. & 
Kew Gard. Misc. 3 (1851) 302; Bot. Voy. Herald 
(1854) 182, t. 32; Steen. Thesis (1927) 1008; Bull. 
Jard. Bot. Btzg III, 10 (1928) 272; Back. & Bakh. 
/. Fl. Java 2 (1965) 542. 

Tree to 7 m, 20 cm 0. Leaflets oblong, acumi- 
nate, 4-8 by Vii-Vjicm. Flowers cauliflorous, 
nocturnal, white, slightly fragrant. Calyx spatha- 
ceous, 3 cm; corolla 5-6 cm. Berry pale yellow, 
pendent, candle-like, smooth, 30-100 by \ l l 2 - 
2V 2 cm. 

Distr. Panama, cultivated in many tropical 
countries, and in many parts of Malesia in parks 
and gardens. With its waxy-fleshy, candle-like 
fruits a showy plant. 



Vern. Candle tree, E, kaarsenboom, D. 

The flowers are bat-pollinated, as in Crescentia. 
In Panama fruits are utilized as cattle food (See- 
mann). 

Phryganocydia corymbosa (Vent.) Bur. ex K.Sch. 
in E. & P. Nat. Pfl. Fam. 4, 3b (1894) 224, f. 89H; 
Gentry, Ann. Mo. Bot. Gard. 60 (1973) 905; 
Herklots, Fl. Trop. Climb. (1976) 69, col. pi. 6. — 
Spathodea corymbosa Vent. Choix (1807) t. 40. 

Sometimes pseudostipules. Leaflets (4-) 12- 15 by 
(2-)7-9 cm, triplinerved. Calyx spathaceous. Co- 
rolla lavender to blue purple, with a white mouth. 

Distr. Panama to Brazil, cultivated at Seria 
(Brunei). 

A Brunei specimen (van Niel 3826) identified 
by A. A. Atchley (1973). 

Pithecoctenium cynanchoides DC. Prod. 9 (1845) 
193; Fabris, Rev. Mus. La Plata 9, Bot. n. 49 

(1965) 353, f. 19. 

Leaves reniform-triangular acuminate, ciliate at 
the margin, 2 l l 2 -A by 2-4 cm. Corolla 3-6 cm, 
white with yellow markings in the mouth. 

Distr. Southern Brazil to Argentina, in E. Java 
once cultivated (Mt Tengger: Buysman). 

Podranea ricasoliana (Tanf.) Sprague, Fl. Cap. 4, 
2 (1904) 450 ; Sykes, New Zeal. D.S.I. R. Inf. ser. 54 

(1966) 39, f. 16; Gentry, Ann. Mo. Bot. Gard. 60 
(1973) 916, f. 29; Herklots, Fl. Trop. Climb. 
(1976) 70, f. 92. — Tecoma ricasoliana Tanf. Bull. 
Soc. Tos. Ort. (1887) 17, t. 1-2. — Pandorea ricaso- 
liana Baill. Hist. PI. 10 (1891) 40. 

Leaflets 7-9, crenate, 2 l [ 2 -A by 1-2 cm. Corolla 
pale lavender or pinkish, with magenta patches and 
lines in the mouth and tube, 6-8 cm long, in 
terminal thyrses. 

Distr. South Africa, cultivated elsewhere in the 
tropics (e.g. in the Philippines, New Caledonia) and 
subtropics (Mediterranean, New Zealand). 

Pyrostegia venusta (Ker) Miers, Proc. R. Hort. 
Soc. Lond. 3 (1863) 188; Steen. Bull. Jard. Bot. 
Btzg III, 10 (1928) 189; Bruggeman, Ind. Tuinb. 
(1939) 64, f. 21 ; Back. & Bakh./. Fl. Java 2 (1965) 
536; Sykes, New Zeal. D.S.I.R. Inf. ser. 54 (1966) 
22, f. 6; Herklots, Fl. Trop. Climb. (1976) 71, 
f. 94. — Bignonia venusta Ker, Bot. Reg. (1818) 
t. 249; Bot. Mag. (1819) t. 2050. — Tecoma 
venusta Lem. Hort. Univ. (1834) 1, icon. — P. ignea 
(Vell.) Presl, Bot. Bemerk. (1845) 93. 

Leaflets ovate-lanceolate, 4-6 by 3-4 cm. 
Thyrses dense, rachis 10 cm. Calyx 6-7 mm. 
Corolla tube c. 7 cm. 

Distr. Brazil, commonly cultivated throughout 
the tropics and also in subtropics (North I. of New 
Zealand), in the temperate zone in glass-houses 
since the early 19th century. Widely cultivated in 
Malesia. 

A richly flowering ornamental, good for walls 
and trellis, excellent between 500-1700 m altitude, 
not flowering in Java below 250 m and never setting 
fruit. The oranje stephanoot, D, is easily propagated 
by tjankoks (marcotting) and cuttings. 

Saritaea magnifka (Steen.) Dugand, Caldasia 3 
(1945) 263, fig.; Gentry, Ann. Mo. Bot. Gard. 60 
(1973) 920, f. 31; Santisuk, Kew Bull. 28 (1973) 



1977] 



Bignoniaceae (van Steenis) 



185 



184; Thai For. Bull. Bot. 8 (1974) 46; Herklots, 
Fl. Trop. Climb. (1976) 71, f. 95. — Arrabidaea 
magnified Steen. Rec. Trav. Bot. Need. 24 (1927) 
830, excl. svn. Bignonia magnified Bull; Bull. Jard. 
Bot. Btzg III, 10 (1928) 191; Guttenberg, Ann. 
Jard. Bot. Btzg 44 (1934) 195; Chatterjee, Bull. 
Bot. Soc. Beng. 2 (1948) 78; Back. & Bakh./. Fl. 
Java 2 (1965) 536. — Arrabidaea sp. : Daubanton, 
Teysmannia 29 (1918) 51-53, phot. 

Pseudostipules present. Tendrils simple. Leaflets 
obovate, bluntly short-acuminate, 5-1 1 by 3-6 cm. 
Calyx cupular-campanulate, truncate, 6-12 by 
3-7 mm. Corolla purple to magenta, 8-9 cm, the 
mouth white with magenta markings. 

Distr. Colombia and Ecuador, widely culti- 
vated through the tropics and common in SE. 
Asia and Malesia, never setting fruit. Obviously 
first introduced in Singapore; the Bogor Botanic 
Gardens received it from Banka in 1911. 

A robust climber, good for walls and trellis, 
profusely flowering all the year round, especially 
in the wet season (Oct. -May), easily propagated 
by cuttings, found up to c. 1000 m. 

Spathodea campanulata P. Beauv. Fl. Oware 
Benin 1 (1805) 47, t. 27; Hook. Bot. Mag. 85 
(1859) t. 5091 ; Sprague, Fl. Trop. Afr. 4, 2 (1906) 
529; Merr. Fl. Manila (1912) 429; En. Philip. 3 
(1923) 447; Steen. Rec. Trav. Bot. Need. 24 
(1927) 945; Bull. Jard. Bot. Btzg III, 10 (1928) 232; 
Holttum, M.A.H.A. Mag. 3 (1933) 186, fig.; 
Irvine, Woody PI. Ghana (1961) 739, t. 7; Heine, 
Fl. Trop. W. Afr. ed. 2, 2 (1963) 386; Back. & 
Bakh./. Fl. Java 2 (1965) 540. 

Large tree, 7-25 m, 10-50 cm 0. Pseudostipules 
leafy. Leaves decussate; leaflets in (4-)5-6(-9) 
pairs, oblong, entire, glabrous or puberulous 
beneath, 5-14 by 2 x l 1 -5 l j 1 cm. Flowers erect, in 
terminal thyrses; calyx navicular, ribbed, beaked, 
thinly velutinous, 4-7 cm ; corolla scarlet to orange, 
wide, 8-14 cm. Capsules erect, lanceolate-oblong, 
15-20 by 2 1 2 -3cm; valves keeled; seeds 2'/ 2 by 
2 cm, hyaline winged. 

Distr. Tropical Africa; widely cultivated in the 
tropics in parks and as an avenue tree; throughout 
Malesia, up to c. 1000 m. 

Vern. African tulip, Tulip tree, E, spuitjesboom, 
D, panchot, Malaya, djati bilanda, Kangean; 
Sabah: anchit anchit, maundi, Banggi. 

The flowers are frequented by birds on which 
Beumee (Trop. Natuur 14, 1925, 28-30, f. I) did 
observations at Bogor. Flowers remain open for 
at least two days and each flower may be visited by 
more than one bird, obviously in search of honey. 
Possibly the birds play a role in pollination; 
corollas are frequently damaged. 

•\ ■ i vsu observed bats visiting the flowers at 
night (Ann. Mo. Bot. Gard. 61, 1974, 713). 

The calyx, which is closed in bud, contains water 
and children play the 'waterspout' with it, hence 
the Dutch name. Children also use the boat-shaped 
tor making small sailing boats. 

1 ruiting and flowering lakes place throughout 
the year. At Bogor I found young sccdlr. 
hedges and gardens, with a long taproot .mil dentate 
but there are no records of spontaneous 
naturalization, limber is weak and worthless. 

Kii.n ■ n I. Mai. Pea 2, i'>2}, 547) mentioned 

it erroneously from Singapore under the name 



S. nilotica Seem. ; although this is a closely related 
African tree, it is a distinct taxon characterized by a 
longer-tomentose, hardly ribbed calyx, leaflets 
tomentose underneath, a longer lobed disk and a 
long-hairy ovary. 

Tabebuia chrysantha (Jacq). Nichols. Diet. Gard. 
4 (1887) 1; Gentry, Ann. Mo. Bot. Gard. 60 
(1973) 941. — Bignonia chrysantha Jacq. Hort. 
Schoenbr. 2(1797)45, t. 211. 

Deciduous tree to 25 m, 50 cm 0. Leaflets 
elliptic-oblong, abruptly acuminate, 5-25 by 4-1 1 
cm. Calyx shallowly lobed, 5-13 mm. Corolla 
glabrous, tubular-infundibuliform, 5-8 cm. 

Distr. Mexico to northern Venezuela, in SE. 
Asia cultivated, rare in Malesia (Philippines: U.S. 
Cemetery, Fort McKinley, Rizal Prov.). No fruit 
seen. 

A yellow-flowered species has been reported by 
Back. & Bakh. /. (Fl. Java 2, 1965, 539) to be 
cultivated in Java under the name T. capitata 
(Bur. & K.Sch.) Sandw., but in absence of 
material I cannot check the identity. 

Tabebuia pallida (Lindl.) Miers, Proc. R. Hort. 
Soc. 3 (1863) 199; Gentry, Ann. Mo. Bot. Gard. 
60 (1973) 950. — Bignonia pallida Lindl. Bot. Reg. 
(1826) 12, t. 695. 

Shrub or small tree. Leaves 1-5-foliolate, leaflets 
elliptic to elliptic oblong or obovate, obtuse. 
Inflorescence few-flowered, often reduced to 1 or 2 
flowers. Corolla pinkish lavender to almost white, 
the throat opening yellow. 

Distr. A common variable West Indian species, 
closely related to T. rosea, cultivated in various 
parts of the tropics, in Indonesia distributed from 
Botanic Gardens, Bogor. 

Tabebuia rosea (Bertol.) DC. Prod. 9 (1845) 215; 
Sandw. Kew Bull. (1953) 454; Gentry, Ann. Mo. 
Bot. Gard. 60 (1973) 951, with synonymy. — 
Tecoma rosea Bertol. Fl. Guat. (1840) 25. 

Deciduous tree, up to 30 m, 1 m 0. Leaflets and 
petiolules often unequal, lepidote, 5-30 by 2-12 cm. 
Calyx cupular, bilabiate, densely lepidote, 3 / 4 -2 cm. 
Corolla outside glabrous, 6-10 cm. 

Distr. S. Mexico to Venezuela, cultivated in the 
tropics in parks and along roadsides; a magnificent 
ornamental when abundantly flowering with 
blossoms in bunches on the bare twigs; not rare in 
SE. Asia, in Malesia only known to me from the 
vicinity of Manila, Luzon. No fruit seen. 

Tecoma capensis (Thunb.) Lindl. Bot. Reg. 13 
(IN2K) t. 1117. — Bignonia capensis Thunb. Prod. 
(1800) 105. — Tecomaria capensis (Thunb.) Spach, 
Hist. Nat. Veg. 9 (1840) 137; Sprague, Fl. Cap. 
4, 2 (1904) 448; Steen. Thesis (1927) 832; Bull. 
Jard. Hot. Btzg III, 10 (1928) 193; Bruggiman, 
Ind. Tuinb. (1948) 39, 268 f. 273; BACK. & Baku. 
/. II. Java 2 (1965) 538; Bkummiii. Bull. Jard. 
Bot. Nat. Belg. 44 (1974) 421, f. I (map); Hikk- 
iois, II. Trop. (limb. (1976) 74, f. 101. 

An eiect ami scrambling shrub, up to e. 3 m. 

never in my experience a true climber. I callus 2 1 

pan., crenate, with domatia. I 3 by I 2cm. 
Corolla tubular, red, rarely pale yellow, 4 *> cm. 
Distr. South Africa, cultivated ami naturalized 

m many subtropical and tropical COUntriC 



186 



Flora Malesiana 



[ser. I, vol. 8 2 



the Mediterranean; also in South America. 
Commonly cultivated in Java and elsewhere in 
Malesia, up to c. 1000 m. Flowering all the year 
round. No capsules seen; not run wild. Easily 
propagated by suckers or cuttings. A nice ornamen- 
tal for gardens and often used for hedges. 

Brummitt I.e. assumes that there is only one 
Tecoma (Jecomaria) in Africa and reduced 7 other 
names, distinguishing the tropical taxa as a separate 
race, ssp. nyassae (Oliv.) Brummitt. 

In South African parks and gardens is also 
cultivated an even sulphur-yellow variety which was 
originally found in the wild, cf. A. Jacot-Guil- 
larmod, Veld en Flora 4 (1974) 36; it hybridizes 
with the red-flowered variety and sets abundant 
fruit. 

Tecoma capensis, the Cape Honey suckle, is bird- 
pollinated; cf. M. S. Evans, Nature 18 (1878) 543; 
Scott-Elliot, Ann. Bot. 4 (1890) 270. 



Tecoma stans (L.) H.B.K. Nov. Gen. Sp. 3 (1819) 
144; Corner, Wayside Trees (1940) 170, f. 44, 
pi. 159; Back. & Bakh./. Fl. Java 2 (1965) 539. — 
Stenolobium stans (L.) Seem. Ann. Mag. Nat. Hist. 
10(1862) 30. — See for elaborate treatment p. 135. 

Yellow Bells is a small, erect, ornamental shrub 
with showy yellow flowers. 

Distr. Southern U.S.A. to southern Argentina, 
commonly cultivated through the tropics, in cer- 
tain places naturalized. Easily propagated by seed 
or by suckers or cuttings. It flowers and fruits 
profusely in Malesia the year round, from the 
lowland up to c. 1000 m. Suitable for gardens and 
parks. 

There is a form with hairy leaves (var. velutina 
DC. = T. mollis H.B.K.) and one with much 
incised leaves {var. incisa G. Don), the latter being 
naturalized in Malesia. 



CRYPTERONIACEAE (R. J. van Beusekom-Osinga, Leyden) 

Evergreen trees. Twigs terete to quadrangular, the younger ones mostly with four 
narrow ribs or wings, with thickened nodes, petiole-bases mostly connected by a 
faint line. Leaves with minute or rudimentary stipules, opposite, simple, entire, 
penninerved, shortly petioled, with arched or almost straight nerves mostly anasto- 
mosing in a marginal vein. Inflorescence terminal or axillary, sometimes below the 
leaves, paniculate, copiously branched to extremely depauperate, branched up to 
the third order, with decussate side axes which are sometimes arranged (sub)- 
verticillately or subumbellately by contraction, either ending in profuse to very 
poor racemules, or in cymoid florescences. Bracts mostly small to minute, some- 
times with rudimentary stipules. Flowers (very) small, shortly pedicelled, bisexual, 
sometimes by reduction unisexual and then trees dioecious, actinomorphous, peri- 
to epigynous, (4-)5(-6)-isomerous, sometimes with twice the number of stamens; 
receptacle widely campanulate. Sepals valvate, triangular to deltoid, mostly per- 
sistent. Petals more or less rudimentary, sometimes absent, alternisepalous, inserted 
on the margin of the receptacle, inflexed and enveloping the stamens, valvate, 
rarely imbricate, sometimes cohering, soon caducous. Stamens if isomerous epi- 
petalous, (alternisepalous), inserted on the margin of the receptacle, inflexed in bud, 
persistent or caducous; filaments sometimes very short; connective wide, with or 
without a tendency to conduplication, or completely conduplicate, sometimes with 
a dorsal tubercle or a large outgrowth; anthers adnate, marginal or submarginal, 
linear to semiorbicular, lengthwise dehiscent, introrse to latrorse. Ovary superior or 
inferior, 2-4(-5)-carpellate, 1-6-locular, septs not, or rarely partly, connate; style 
terminal, rather long to short, ± terete, mostly persistent; stigma capitate or 
punctate. Ovules situated in horizontal or vertical position, 1, 2, 3, or many per 
locule, anatropous; placentation parietal, septal, or basal. Fruit superior or 1 / 2 - or 
3 / 4 -inferior, a chartaceous or woody capsule, subglobose to ellipsoid, small to big, 
loculicidally dehiscent with 2-6 valves, on the top often with the persistent style and 
stigma. Seeds few or many, flat, usually small, depressed-ellipsoid, situated basally, 
apically, centrally, or laterally in its membranous wing in which the raphe is 
running freely; endosperm none; embryo straight. 



Distribution. Pantropical, 5 genera and 11 spp., 3 genera in Indo-Malesia (of which one 
endemic in Borneo), one monotypic genus in S. Africa (Rhynchocalyx) and one in Peru and 
Bolivia (Alzatea). 

Ecology. Lowland and hill tropical forest, up to 1300 (-1700) m, mostly in rain-forest, but 
Crypteronia paniculata not shunning a more or less seasonal climate. 

Dactylocladus stenostachys is a characteristic peat swamp forest tree. 

Young branches of Crypteronia macrophylla are frequently inhabited by ants. 

Young foliage seems often bright coloured: in Axinandra innovations are mauve, while in 
Crypteronia paniculata young leaves and twigs are deep blue or violet turning pinkish brown 
then green (C'ornf.r), in other species they are purplish. 

Morphology. In all genera the nodes of the twigs are thickened and a characteristic trans- 
versal ridge or line, sometimes faint (absent in Rhynchocalyx), connects the leaf-bases. On the 
intcrnodes four lengthwise raised lines or narrow wings occur, especially distinct in the upper 
part of young twigs; they wear off later. 

The leaves olfcr no significant characters, but interesting is the occurrence of ephemerous 
rudimentary stipules which can only be observed on innovations. These are common among 

(187) 



188 Flora Malesiana [ser. I, vol. 8 2 

Myrtalean families, but obviously absent in Melastomataceae, where these structures were not 
found in a sampling of fifteen genera. 

The petals in Crypteroniaceae are reduced to a varying degree or are even totally absent (in 
Crypteronia). Moreover, they are always soon caducous, except in Alzatea where they are almost 
invisible and mucilaginous. In all genera they are conduplicate and enveloping the inflexed 
stamens as a hood. In Dactylocladus and Rhynchocalyx they are minute and unguiculate; in 
Axinandra they are proportionally bigger, and have a broad instead of an unguiculate base. The 
petals of Axinandra are complicated and show a highly interesting specialization; they are 
coherent to connate, together having the shape of an umbrella or a mushroom. Their wide, 
tapering basal parts together form the awning of the umbrella, the narrow, coherent, median 
parts form the stem of the umbrella, and often there are wider, frayed, reflexed, apical parts of 
the petals together forming the handle of the umbrella (or the 'root' of the mushroom). This 
whole structure envelops the stamens very closely, and drops when the flower opens and the 
inflexed stamens stretch. It is, furthermore, interesting that these petals of Axinandra, depending 
on the species, can be valvate-connate, valvate- (or somewhat imbricate-)conduplicate, or imbri- 
cate-contorted. 

Several petal characters of Crypteroniaceae are found again scattered in other Myrtalean 
families. Reduction, absence, as well as caducousness of petals occurs sporadically in almost 
all of these families. Unguiculate petals are more or less characteristic for Lythraceae, but are 
also found in Sonneratiaceae (Duabanga), and in Rhizophoraceae {e.g. Carallia). Connate petals 
with a broad base, sometimes fused to a 'cap' occur in Myrtaceae. Coherence of petals is also 
present in Rhizophoraceae (Ceriops). Valvate and imbricate petals, both found in one genus, 
Axinandra, are usually family characters in Myrtales. Contorted petals are, apart from Axinandra, 
only found in Melastomataceae. 

The enveloping of the stamens by the petals in all petal-bearing Crypteroniaceous genera is 
almost unique in Myrtales, being only found in a few Rhizophoraceae (Rhizophora and Bruguiera). 
However, in the latter family the petals do not cover the stamens as a hood, as is the case in 
Crypteroniaceae. This is one of the characters upon which the identity of Crypteroniaceae as a 
family is based. 

The stamens in Crypteroniaceae are arranged in one isomerous epipetalous (alternisepalous) 
whorl, except in Axinandra, where one diplostemonous whorl is present. In general number and 
position of the stamens in Myrtales can be derived from a situation with two isomerous whorls, 
either by reduction or by polymerisation ('dedoublement') and multiplication. Arrangement of 
stamens in two isomerous whorls, the diplostemonous androecium, is mostly considered to 
represent the basic structure of the androecium in Myrtales. Melchior (in Engl. Syllabus 2, 1964, 
345) distinguished for the androecium in Myrtales two progressive trends both starting from the 
diplostemonous androecium, viz multiplication into many stamens in many whorls and reduction 
towards the haplostemonous state and even eventually to 3, 2, or 1 stamen(s). 

In all Crypteroniaceous genera the stamens are inflexed in bud. This is a widespread character 
in Myrtales, being rather typical for this order. In some families {Myrtaceae, Rhizophoraceae) it 
is not present in all genera, and Lythraceae, Onagraceae, and Haloragidaceae are the only families 
in which it is totally absent. The total absence of inflexed stamens in the Lythraceae is another 
fact which militates against inclusion of Crypteroniaceous genera in that family. 

The gynoecia of the Crypteroniaceous genera are distinguished by cells which are divided by 
interrupted septs, though these may touch each other in the centre of the gynoecium. This 
(hemi)synplicate condition is very rare and assumed to be primitive within the Myrtales, in which 
it is only found in Crypteroniaceae and in a few genera of Myrtaceae. 

The capsules of Crypteronia and Axinandra show interesting specialized structures with a func- 
tional significance with regard to opening and closing of the capsule. The mechanisms for this are 
based upon hygroscopical properties of fibres in vascular bundles. 

The morphology of the seed in the Crypteroniaceae is peculiar. The seed-coat forms a flat, 
membranous wing, through which the raphe is running from the insertion to the top where it 
usually takes a more or less sharp turn, and runs back towards the seed proper, which either 
takes a central, apical, lateral, or basal position in the wing. This is another assumedly primitive 
character within Myrtales, again only found in Crypteroniaceae and in a few genera of Myrtaceae. 



1977] Crypteroniaceae (van Beusekom-Osinga) 189 

This character is also rare in other orders. It was first discovered in the Trochodendraceae and is, 
therefore, indicated by me as the Trochodendraceous seed type. 

Summarizing, we find that almost the whole variety of floral characters in the Crypteroniaceae 
is also found scattered in other Myrtalean families. In this respect the Crypteroniaceae are rather 
heterogeneous, though not more than for instance the Myrtaceae and the Melastomataceae. On 
the other hand, the family is unique in the Myrtales by having petals enveloping the stamens as a 
hood. Moreover, the presence of one whorl of epipetalous (alternisepalous) stamens, characteris- 
tic for four out of the five Crypteroniaceous genera, is very rare in other Myrtalean families, 
being restricted to one or two genera of the Myrtaceae and of the Lythraceae, and to the mono- 
typic Oliniaceae. Finally, the conduplication of the connective or the tendency to it in all Crypter- 
oniaceae except Dactylocladus, is another important family character, in other Myrtales only 
found in a few Melastomataceae. Apart from the above-mentioned characters Crypteroniaceae 
are also characterized by the septation of the gynoecium and by the Trochodendraceous seed- 
structure, both being only found in other Myrtales in a few Myrtaceous genera. They are, how- 
ever, from a practical viewpoint, less useful for easy diagnosis. — C. F. van Beusekom. 

Taxonomy. Crypteroniaceae belong undoubtedly to Myrtales. Though the family concept in 
this order is fairly satisfactory, it can be observed from the above-made remarks that there are 
not many exclusive characters, most of them breaking down occasionally in one family, or 
occurring also sporadically in another family. Each family in Myrtales seems to be characterized 
by a unique character combination in addition to one or two exclusive characters. 

For Crypteroniaceae this combination and characters are: swollen nodes with transversal line, 
internodes with lengthwise raised lines or wings, petals in bud hood-like enveloping the stamens, 
soon caducous (in Crypteronia absent), stamens inflexed in bud and in one epipetalous whorl 
(except in Axinandra in two whorls), absence of a perianth tube or of any space between the 
insertion of petals and stamens, and furthermore the presence of a (hemi)synplicate gynoecium 
and seeds of the Trochodendraceous or related type, both assumedly primitive characters, and 
almost exclusive within Myrtales. 

Palynological evidence does not fully sustain the recognition of Crypteroniaceae as a distinct 
family; it could be accepted, but the evidence may allow other possibilities (Muller, Blumea 22, 
1975, 275). 

Anatomical evidence is not much in favour of the family concept as proposed; the genera could 
in this respect be divided up among Melastomataceae and Lythraceae, but it remains to be seen 
in how far anatomical characters clearly sustain other current family concepts in Myrtales. 

Myrtales are certainly a very ancient complex and during their evolution advanced characters 
have evolved, reduction series occurred in more lines, and primitive characters may have inciden- 
tally persisted in various branchings of ancestral tree in taxa which are not necessarily viewed as 
closely related. This would also explain the 'reticulate character distribution', a condition found 
in several families of Myrtales. 

None of the Myrtalean families is really homogeneous, but from this can and should not be 
concluded that these families are unnatural. For tracing ancestry and evolution naturalness is 
more important than homogeneity which, properly, increases always with decreasing taxonomic 
rank. 

The main subdivision of two subfamilies is supported by wood-anatomical characters (van 
Vliet, J. Micr. 104, 1975, 65), but other anatomical characters (van Vliet & Baas, Blumea 22, 
1975, 175) and palynological data (Muller, I.e.) do neither support it, nor militate against it. At 
tribal level morphological, anatomical and palynological data appear not to agree. The sub- 
division adopted here is based on morphology. — C. F. van Bum kom. 

Anatomy. Van Vi.m.t, J. Microscopy 104 (1975) 65 82 (wood anatomy and relationships); 
van Vliet & Baas, Blumea 22 (1975) 175 195 (leaf, nodal and twig anatomy). These two papers 
contain a full bibliography to the older literature. 

The anatomy of Crypteroniaceae term ktto is heterogeneous. Distinctive characters are: 
cuticle granular (Axinandra, Crypteronia) Of smooth (Dactylocladus)', stoma ta pa racy tic (A.xinan- 

dra, Crypteronia) or anomocytic (Dactylocladus)', hypodennii present (Axinandra. Crypteronia) 

or absent (Dact) locladus)', petiole with arc-shaped vascular bundle (.1 \mandra) Of With I closed 

system (Crypteronia, Dactylocladus)', phloem with styloid crystals (Axinandra, Crypteronia) or 



190 Flora Malesiana [ser. I, vol. 8 2 

crystal sand (Dactylocladus); cork arising in pericycle (Axinandra, Crypteronia) or subepidermal 
(Dactylocladus); node complex, with complete girdling trace (Axinandra, Crypteronia 
paniculata) or with common gaps (other species of Crypteronia, Dactylocladus); cortical 
bundles present (Axinandra p.p.) or absent (other taxa); vesturing of vessel pits in wood confined 
to the pit chamber (Axinandra, Dactylocladus) or also on pit apertures (Crypteronia); vessel-ray 
pits alternate (Axinandra, Crypteronia) or reticulate to scalariform and larger (Dactylocladus); 
parenchyma aliform with narrow wings (Axinandra, Dactylocladus) or chiefly diffuse in aggregates 
(Crypteronia); rays heterogeneous Kribs type I (Axinandra, Crypteronia) or Kribs type III 
(Dactylocladus); intercellular canal-like spaces present in rays (Crypteronia, Dactylocladus) or 
absent (Axinandra). 

The entire evidence from vegetative anatomy supports affinities between Crypteronia and 
Axinandra — both genera sharing salient features with a number of Melastomataceae. Dactylo- 
cladus resembles several Melastomataceae in its anatomy more closely than it does Axinandra or 
Crypteronia. The inclusion of Alzatea from S. America and Rhynchocalyx from S. Africa in the 
Crypteroniaceae adds to the anatomical heterogeneity of the family. On anatomical grounds, 
Rhynchocalyx fits better in Lythraceae, and Alzatea could also be accommodated in that family 
with its trilacunar nodes as only aberrant character. The existence of a considerable overlap of 
the anatomical range in Melastomataceae with that of Lythraceae, Sonne r at iaceae and Oliniaceae, 
forbids, however, formal taxonomic decisions on anatomical grounds only. — P. Baas. 

Palynology. Pollen grains are small, ranging in size from 11 \im in Crypteronia paniculata to 
20 urn in Rhynchocalyx lawsonioides, and thin-walled with a smooth or finely verrucate outer 
surface. In Alzatea they are tricolporate, in Axinandra, Dactylocladus and Rhynchocalyx hetero- 
colpate, while Crypteronia is characterized by bilaterally flattened bisyncolporate grains (Muller, 
1975). 

The subdivision of the family according to the pollen types is not correlated with those based 
on morphological or anatomical characters. 

The relatively unspecialized Alzatea type occurs in many dicotyledonous families. The hetero- 
colpate type is found in Combretaceae, Lythraceae, Melastomataceae, Oliniaceae, and Penaea- 
ceae. Pollen grains similar to the Crypteronia type only occur in Cunoniaceae, but differ in 
sculpture and an affinity with this family, as suggested by Erdtman (1952), appears remote. — 
References : Erdtman, Pollen morphology and plant taxonomy. Angiosperms. Stockholm (1952); 
Muller, Note on the pollenmorphology of Crypteroniaceae s.l. Blumea 22 (1975) 275. — J. 
Muller. 

Uses. Crypteronia paniculata, which may attain a height of 30 m, is said to have durable, 
reddish heartwood and is sometimes used in West Java for house-building; also in S. Sumatra 
reports are favourable (Heyne, Nutt. PI. 1927, 1158), but occurrence is too scattered to have 
come into general use. 

Dactylocladus stenostachys is one of the most important export timber trees of Sarawak and 
Sabah; see under that species. 



KEY TO THE GENERA 

1. Flowers sustained by 1 bract. Stamens as many as sepals; connective not or only slightly conduplicate. 

Capsule small, chartaceous. Seed situated (latero-)centrally in its wing. Tribe Crypteronieae. 
2. Petals absent. Ovary superior with many ovules per cell. Capsule with many seeds per cell. Nerves 

distinct, ascending and often anastomosing into a looped marginal nerve 1. Crypteronia 

2. Petals present, soon caducous. Ovary at least half-inferior, the lower part immersed in the receptacle, 
with 3 ovules per cell. Capsule with 1-3 seeds per cell. Leaves coriaceous with vague venation 

2. Dactylocladus 

1. Flowers sustained by 3 bracts, the two lateral ones often minute. Stamens twice as many as petals. 

Connective completely conduplicate. Ovary inferior, with 1 or 2 ovules per cell. Capsule large, woody, 

half-inferior. Seed situated basally in its wing. Nerves ascending and anastomosing into a looped 

marginal nerve. Tribe Axinandreae 3. Axinandra 



1977] Crypteroniaceae (van Beusekom-Osinga) 191 

1. CRYPTERONIA 

Bl. Bijdr. (1826) 1151; Hassk. Cat. Hort. Bog. (1844) 232 {'Crypterhonia'); Bl. 
Mus. Bot. Lugd. Bat. 2 (1856) 123, t. 42; B. & H. Gen. PI. 1 (1867) 782; DC. Prod. 
16, 2 (1868) 677; Clarke in Hook./. Fl. Br. Ind. 2 (1879) 573; Koehne, Verh. Bot. 
Ver. Brandenburg 22 (1881) 69; O. K. Rev. Gen. PI. 1 (1890) 250 ('Crvptoneria); 
Niedenzu, Bot. Jahrb. 15 (1892) 161 ; in E. & P. Nat. Pfl. Fam. 3, 7 (1892) 21, t. 8; 
Hall./. Abh. Naturw. Ver. Hamb. 18 (1903) 90; Med. Rijksherb. 1 (1911) 31; 
ibid. 35 (1918) 17; Hutch. Gen. Fl. PI. 2 (1967) 33 ; Beus.-Osinga & Beus. Blumea 
22 (1975) 258. — Henslouia Wall. PI. As. Rar. 3 (1831) 13, t. 221, non Bl. 1850. — 
Quilamum Blanco, Fl. Filip. 1 (1837) 851 ; ed. 2, 1 (1845) 136; ed. 3, 1 (1877) 245; 
cf. Merr. Sp. Blanc. (1918) 282. — Fig. 1, 2, 6. 

Leaves elliptic or ovate to (ovate-)lanceolate, glabrous or slightly pubescent; 
midrib flat or slightly impressed above, prominent beneath; nerves ascending and 
often anastomosing in a looped marginal nerve, flat above, ± prominent beneath, 
intramarginal nerve mostly present. Panicles terminal or axillary, sometimes on 
leafless older nodes or ramiflorous, erect to usually pendulous, poorly to rather 
copiously branched; axes terete to more or less angular, puberulous; racemules 
with very numerous flowers. Flower-bracts persistent. Flowers bisexual or by 
reduction unisexual and then trees dioecious, 4-5(-6)-isomerous, pedicelled. 
Receptacle in- and outside puberulous, inside sometimes minutely tomentose, 
hardly or not accrescent. Sepals deltoid to triangular, persistent. Petals absent. 
Stamens persistent, in 9 flowers staminodial and mostly permanently inflexed; 
filaments filiform, somewhat flattened, connective about orbicular, with or without 
a tendency to conduplication, dark when dry, anthers apically or laterally on the 
connective, semiorbicular to broad-linear, latrorse or ± introrse. Ovary superior 
or almost so, the lower part adhering to the receptacle, (sub)globose to pyramidal, 
2-4-carpellate, 2-4-celled, with free or only basally connate septs, badly developed 
in j flowers; style filiform to subulate, somewhat longer to shorter than the ovary, 
more or less puberulous, persistent; stigma punctate to capitate. Ovules many, 
either in horizontal position on the septs or in ± vertical position basally between 
the septs. Capsule superior or almost so, (sub)globose or more or less (ob)ovoid, 
puberulous, upper part dehiscent with 2-4 valves, inside split as far as the basal 
connation of the septs; valves at the top kept together by the non-dehiscent part of 
style and stigma. Seeds many, very small, in horizontal or vertical position; seed 
ovoid-ellipsoid, situated latero-centrally in its narrow, membranous wing, which 
has a shorter or longer apical and basal extension, raphe running closely along the 
embryo (microscopical!). 

Distr. 4 spp., of which one ranges through tropical SE. Asia (Assam, Bengal, Lower Burma, Thailand, 
and Indo-C'hina) to Malesia, the other three endemic in Malesia. Fig. 3. 

Ecol. Lowland and montane rain-forests below c. 13(H) m; C. paniculate/ also rather frequent in areas 
with a more or less seasonal climate. Fl. fr. in almost all taxa Jan. Dec. 

Uses. C. paniculalu seems to yield a fairly good timber, but is never found in quantity. 

k i -, i « > rm s c i < 1 1 s 

I Ovary (and capsule) 2-ccllcd with s\5 ovules rq. seeds on the septs. Dioecious tree: flowers by reduction 
unisexual, rarely bisexual I Ipyraceous. Sect Crvpteronia I. C. paniculata 

I Ovary (and capsule) 1- or 4-ccllcd with re ovules (seeds) inserted basally. Flowers bisexual. Leaves 
usually coriaceous. Sect, liauspcrmia. 



192 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 1. Crypteronia macrophylla Beus.-Osinga. a. Inflorescence, x l / 2 , b. flower, x 10, c. stamen, x 15, 

d. fruit, x 10 (Ashton S 19372). 



1977] Crypteroniaceae (van Beusekom-Osinga) 193 



Fig. 2. Crypieronia macrophylla Beus.-Osinga. Young twig with leaves, X '/i (Ashton S 19372). 



194 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 3. Range of the genus Crypteronia Bl. ; sect. Crypteronia unbroken line, sect. Basispermia broken line. 

2. Inflorescences terminal at main or side twigs, 10-25(-40) cm. Ovary (capsule) 3-locular 1 . Sepals 

usually with longitudinal crest at the inside 2. C. cumingii 

2. Inflorescences axillary below the leaves on older nodes, or ramiflorous, 15-90 cm. Ovary (capsule) 
3- or 4-celled. Sepals with or without longitudinal crest at the inside. 
3. Inflorescences below the leaves on older nodes, 15-50 cm, only with primary, very rarely with a few 
secondary side-axes near the base. Ovary (capsule) 3-celled. Sepals without crest . . 3. C. griffithii 
3. Inflorescences ramiflorous, 45-90 cm 2 , with primary, secondary and tertiary side-axes. Ovary (cap- 
sule) 4-celled. Sepals with a longitudinal crest at the inside 4. C. macrophylla 



1. Crypteronia paniculata Bl. Bijdr. (1826) 1151; 
Mus. Bot. Lugd. Bat. 2 (1856) 123, t. 42; DC. Prod. 
16, 2 (1868) 679; Kurz, J. As. Soc. Beng. 46, ii 
(1877) 86, incl. var. glabra (Wall.) Kurz et var. 
pubescens (Wall.) Kurz; Fl. Burma 1 (1877) 
519; Clarke, Fl. Br. Ind. 2 (1879) 574; Niedenzu, 
Bot. Jahrb. 15 (1892) 177; K. & V. Bijdr. 1 (1894) 
203, incl. var. leptostachya (Planch.) K. & V.; 
King, J. As. Soc. Beng. 67, ii (1898) 5; Ridl. Agr. 
Bull. Str. & F. M. S. n.s. 1 (1902) 180 ('Crypto- 
rania , )\ Merr. Philip. J. Sc. 1 (1906) Suppl. 102; 
Brandis, Ind. Trees (1906) 341 ; Koord. Exk. Fl. 
Java 2 (1912) 664; Atlas 2 (1914) 318; Merr. Sp. 
Blanc. (1918) 282; Gagnep. & Guillaumin, Fl. 
Gen. I.-C. 2 (1920) 696, t. 70; Ridl. Fl. Mai. Pen. 1 
(1922) 821 ; Merr. En. Philip. 3 (1923) 140; Craib, 
Fl. Siam. En. 1 (1931) 729; Ochse & Bakh. Ind. 
Groent. (1931) 182, f. Ill; Burk. Diet. 1 (1935) 
693; Corner, Ways. Trees (1940) 197, pi. 48, f. 51 ; 
Kraemer, Trees W. Pacif. Region (1951) 330; 
Lecompte, Fl. Camb. Laos & Vietn. 4 (1965) 57; 
Beus.-Osinga & Beus. Blumea 22 (1975) 259. — 
Henslowia glabra Wall. PI. As. Rar. 3 (1831) 14; 



Cat. (1831-32) n. 4903; Planch. Hook. Lond. J. 
Bot. 4 (1845) 478; Miq. Fl. Ind. Bat. 1, 1 (1856) 
716. — C. glabra (Wall.) Planch, ex Endl. Gen. 
PI. Suppl. 4, 2 (1847) 39 ; Bl. Mus. Bot. Lugd. Bat. 2 
(1856) 123; DC. Prod. 16, 2 (1868) 678; Clarke, 
Fl. Br. Ind. 2 (1879) 574; F.-Vill. Nov. App. 
(1880) 91; Vidal, Sinopsis (1883) 27, t. 52, f. E; 
Phan. Cuming. (1885) 48; Rev. PI. Vase. Filip. 
(1886) 138. — Henslowia pubescens Wall. PI. As. 
Rar. 3 (1831) 14, t. 221; Cat. (1831-32) n. 4904; 
Planch. Hook. Lond. J. Bot. 4 (1845) 477, t. 
14 B, f. 1-4; Griff. Not. 4 (1854) 404; Ic. PI. As. 4 
(1854) t. 562 f. 3, t. 564 f. 2; Miq. Fl. Ind. Bat. 1, 1 
(1856) 716. — Quilamum luteum Blanco, Fl. Filip. 
1 (1837) 851; ed. 2, 1 (1845) 136; ed. 3, 1 (1877) 
245; Niedenzu, Bot. Jahrb. 15 (1892) 177. — 
Henslowia affinis Planch. Hook. Lond. J. Bot. 4 
(1845) 477; Miq. Fl. Ind. Bat. 1, 1 (1856) 716. — 
Henslowia leptostachys Planch. Hook. Lond. J. 
Bot. 4 (1845) 478; Miq. Fl. Ind. Bat. 1, 1 (1856) 
716. — Henslowia hookeri Wall. Cat. (1847) n. 
8566, nomen. — C. affinis (Planch.) Planch, ex 
Endl. Gen. PI. Suppl. 4, 2 (1847) 39. — C. 



(1) Rarely a few 4-locular ovaries may be found among the predominant 3-locular ones in one in- 
florescence. 

(2) One should be aware that of the big inflorescences of C. macrophylla usually only fragments have 
been collected. 



1977] 



Crypteroniaceae (van Beusekom-Osinga) 



195 



pubescens (Wall.) Planch, ex. Endl. I.e. ; Bl. 
Mus. Bot. Lugd. Bat. 2 (1856) 123; DC. Prod. 16, 
2 (1868) 678, incl. var. affinis (Planch.) DC; 
Clarke, Fl. Br. Ind. 2 (1879) 574, incl. var. hookeri 
(Wall, ex DC.) Clarke; F.-Vill. Nov. App. 
(1880) 91; Niedenzu, Bot. Jahrb. 15 (1892) 176, 
incl. var. typica. — C. leptostachys (Planch.) 
Planch, ex Endl. Gen. PI. Suppl. 4, 2 (1847) 39; 
Bl. Mus. Bot. Lugd. Bat. 2 (1856) 123; DC. Prod. 
16, 2 (1868) 679; Baill. Hist. PI. 6 (1877) 436, 
f. 414, 415 Cleptostachya'); Vidal, Phan. Cuming. 
(1885) 53; Rev. PI. Vase. Filip. (1886) 139; 
Niedenzu, Bot. Jahrb. 15 (1892) 175; Merr. En. 
Philip. 3 (1923) 140. — Henslowia paniculata (Bl.) 
Miq. Fl. Ind. Bat. 1, 1 (1856) 716. — C. lutea 
(Blanco) Bl. Mus. Bot. Lugd. Bat. 2 (1856) 123; 
DC. Prod. 16, 2 (1868) 679. — C. hookeri Wall, ex 
DC. I.e. — C. wallichii DC. I.e.; Hance, J. Bot. 14 
(1876) 259. 

Tree up to c. 30 m. Twigs glabrous or puberulous. 
Leaves elliptic or oblong or ovate-oblong, (5-)10- 
15(-25) by (3-)5-10(-12)cm, rounded to cuneate 
at the base, acuminate to cuspidate at the top, 
tip usually obtuse, sometimes acute, glabrous or 
puberulous, chartaceous to herbaceous, usually 
finely and distinctly reticulate beneath; nerves 
8-10(-12) pairs; petiole 5-10 mm, 1-2 mm 0, 
glabrous or puberulous. Panicle axillary or termi- 
nal, also on leafless older nodes, (5-)10-30cm, 
branched up to the second order, without definite 
peduncle, more or less pendulous, not woody; 
main axis more or less angular; primary axes 2-5 
pairs, the lower ones usually with 1-3 pairs of 5-25 
cm long racemules. Bracts of axes triangular or 
narrowly triangular, 1-6 mm, usually caducous. 
Bracts of the flowers narrowly triangular to linear, 
V2-I mm, persistent. Pedicels 1-3 mm, c. V 4 mm 0. 
Receptacle c. ' 2 mm high, c. 2 mm wide, puberu- 
lous, inside sometimes minutely tomentose. Sepals 
deltoid to triangular, (" 4 -)' 2 -l'/ 4 mm. Stamens 




Fig. 4. Range of Cryptcroniu paniculata Bl. var. 
paniculata (unbroken line) and far. ttffiutt 
(Plan' 11 j Hi | s.-Osinca (broken line). 



5 (4), reduced and mostly permanently inflexed in 
$ flowers; filaments 2V2-3V2 mm, glabrous; con- 
nective c. l l 2 (- 3 U) mrn, not conduplicate ; anthers 
linear, V4-V2 by c. 1 U mm, latrorse. Ovary 
reduced in <J flowers, subglobose, 2(-3)-locular, 
1-2 mm, puberulous; style 1-2 mm, c. '/4 mm 0, 
puberulous ; stigma capitate, c. 1 / 4 mm. Ovules 
inserted on the surface of the septs. Capsule 
(sub)globose or more or less obovoid, 2-3 mm, 
usually recurved; valves 2, rarely 3. Seeds , / r -l by 
c. 1 U mm; acute apical part of the wing half as long 
as the seed proper, basal part mostly short. 

Distr. Continental SE. Asia (Assam: Khasya 
Hills, Sylhet, Lushai Hills; Bengal; Burma: 
Arakan Div. to Rangoon; S. Indo-China) and 
West Malesia: eastwards to the Philippines, 
Borneo, and Flores. Fig. 4. 

Note. Two replacing varieties are distinguished 
but it should be mentioned that in Penang and in 
the Rangoon area, where the ranges coincide, more 
or less intermediate specimens are found which have 
almost or practically glabrous leaves and twigs. 

KEY TO THE VARIETIES 

1. Leaves and branches glabrous. Receptacle 
inside more or less puberulous. Sepals c. 1 mm 
long. Capsules (sub)globose . var. paniculata 

1. Leaves at least beneath, and young twigs 
puberulous. Receptacle inside tomentose. 
Sepals Vr-ViC-*/^) mm l° n 8- Capsules (sub)- 
globose to obovoid var. aflinis 

var. paniculata. — C. paniculata, incl. var. glabra et 
var. leptostachya — C. glabra — Quilamum luteum 
— C. leptostachys — C. lutea — C. wallichii. 

Leaves (5-)10-15(-25) by (3-)5-10(-12) cm, 
glabrous; petiole glabrous. Buds glabrous. In- 
florescence up to c. 30 cm; racemules up to 25 cm. 
Receptacle inside puberulous. Sepals c . 1 mm long. 
Staminodes V2-I mm long. Capsule (sub)globose. 

Distr. Continental SE. Asia: E. Bengal (Chitta- 
gong), Assam (Khasia Hills, Lushai Hills), 
Andaman Is., Burma (Arakan, Rangoon, Pegu, 
Tenasserim); in Malesia: Malay Peninsula (Kedah, 
Perak, Langkawi I., Pahang, Selangor, incl. 
Penang I.), Sumatra (Sibolangit, Pajakumbuh, 
Painan, Palembang), Java, Lesser Sunda Is. (Bali, 
Sumbawa, Flores), Borneo (Sabah, W. Kutai), 
Philippines. Fig. 4. 

Ecol. Primary and secondary forests on hills 
and mountains up to 1700 m. Reported from river- 
banks, ridges, ravines and forest borders. Scattered, 
locally fairly common. 

Field notes. Bole up to 15 m, often crooked. 
Buttresses sometimes present. Bark smooth or 
rough, scaling off. Young twigs and leaves deep 
blue or violet (Corner), when dried blackish. 
Flowers white, pale green, or greenish yellow. 

Uses. The timber is reported as of good quality, 
used amongst others for house-building purposes 
and cart-wheels. In the Philippines the bark is 
sometimes used against skin-eruptions. In Java the 
young shoots are eaten with rice as lalab (Hass- 
karl). 

Vern. Sumatra: kayu kapax, mtda/lg ajam, M; 
Malaya: bikoi, bikwol, berkol, bua babt, mStucuah, 

ruptii, tukol, M; Java: ktbuiun, kitjrngklak, S, 
kayu babi, kigandik, ponggokan (Jakarta, M), 



196 



Flora Malesiana 



[ser. I, vol. 8 2 



kibakko, kidjarak (Priangan), bang-kongan (Ban- 
ten), blis or blisan, kayu tjeleng (Banjumas), 
mungur (Madiun), wungu lubu (Kediri), tjeleng(an,) 
wrakas or kwakas (Pekalongan), glingsing (Pasu- 
ruan), sepat (Besuki), all J; Flores: madja; Sum- 
bawa: sarowe; Borneo: kinkidon mantok, Dusun; 
Philippines: balinog, bitog, bitok, bongari, bongaui, 
bungaing, kilamo, malabaydnas, salasan, tiaui, tua, 
Tag., agidai, agudai, barabok, barakbak, bungaing, 
Ilk., banujo, tolan-manok, P.Bis., baroga, baruga, 
kodai, kudai, ladao, Ig., kamanok, Bis., malabiong, 
Sbl. 

var. affinis (Planch.) Beus.-Osinga, comb. nov. — 
Henslowia affinis Planch. Hook. Lond. J. Bot. 4 
(1845) 477. — C. affinis — C. pubescens, inch var. 
affinis (Planch.) DC. Prod. 16, 2 (1868) 678. — 
C. hooker i. 

Leaves (2-)5-10(-16) by (lV 2 -)3-6(-8) cm, 
sometimes sparsely puberulous above, more or less 
puberulous beneath especially on the nerves; 
petiole puberulous. Buds puberulous. Inflorescence 
up to c. 20 cm; racemules up to 20 cm. Receptacle 
inside tomentose. Sepals 1 U- 1 I 2 (- 3 U) mm long. 
Staminodes 0.1-0.3 mm long. Capsule (sub)- 
globose to obovoid. 

Distr. Continental SE. Asia: Burma (Rangoon, 
Pegu, Tenasserim Div.), Thailand, Laos, Cam- 
bodia, S. Vietnam; in Malesia: Malaya (Kedah; 
Penang I.). Fig. 4. 

Ecol. Several times reported from riverbanks 
and ridges, 90-300 m. 

2. Crypteronia cumingii (Planch.) Planch, ex 
Endl. Gen. PI. Suppl. 4, 2 (1847) 39; Bl. Mus. 
Bot. Lugd. Bat. 2 (1856) 123; DC. Prod. 16, 2 
(1868) 678; F.-Vill. Nov. App. (1880) 91 ; Vidal, 
Phan. Cuming. (1885) 20; Rev. PI. Vase. Filip. 
(1886) 138; Niedenzu, Bot. Jahrb. 15 (1892) 179; 
Merr. Philip. J. Sc. 1 (1906) Suppl. 102; En. 
Philip. 3 (1923) 139; Merr. & Perry, J. Am. Arb. 
12 (1941) 270. — Henslowia cumingii Planch. 
Hook. Lond. J. Bot. 4 (1845) 478, t. 14, f. C 1-4. — 
C. javanica Baill. Hist. PI. 6 (1877) 435, f. 412, 
413, nomen. — C. laxa Elmer, nomen in sched., cf. 
Merr. En. Philip. 3 (1923) 140. 

Tree up to 40 m. Twigs glabrous. Leaves elliptic 
to ovate-oblong, (5-)8-25 by (3-)4— 14 cm, usually 
emarginate sometimes rounded or acute at the base, 
acute, sometimes acuminate at the top, tip mostly 
obtuse, glabrous, coriaceous, distinctly and slightly 
prominently reticulate beneath; nerves 6— 12(— 15) 
pairs; petiole 2— 10(— 15) mm, 1-4 mm 0, glabrous. 
Inflorescence terminal, at the end of main or side 
twigs, 10— 25(— 35) cm, branched up to the first, 
second or third order, whether or not peduncled, 
erect, with flaccid to stiff and sublignose axes ; main 
axis more or less flattened; primary axes 5-25, 
paired or irregular, sometimes in whorls of 4; 
racemules up to 15 cm. Bracts of axes triangular, 
2-10 mm, sometimes present as small or reduced 
leaves, persistent or caducous. Bracts of flowers 
narrowly triangular, l l 2 -\ l l 2 mm, persistent. Pedi- 
cels lV 2 -2 mm, c. l / 2 mm 0. Receptacle l l 2 -l l U mm 
high, V/i^limm wide, densely whitish papillose 
inside. Sepals 5, triangular, 1— 1 1 / 4 (— 1 1 / 2 ) mm, 
inside mostly with a longitudinal densely puberu- 
lous crest. Stamens 5; filaments 1-2V 2 mm, glab- 
rous; connective c. V4 mm, slightly conduplicated ; 



anthers linear or semiorbicular, c. 1 U(- 1 I 2 ) by 
0.1-0.2 mm, ± latrorse. Ovary subglobose, 3- 
celled, 1-1 */a mm, whitish papillose and/or puberu- 
lous; style (1— )l 1 / 2 -2 1 / 2 mm, 0.2-0.4 mm 0, more 
or less shortly puberulous to papillose; stigma 
punctate to slightly capitate. Ovules inserted on the 
somewhat conically elevated edges between the 
bases of the septs. Capsule (sub)globose, more or 
less episeptally impressed, c. 2 mm; valves 3. 
Seeds c. 3 / 4 by c. 1 U mm, apical and basal part of 
wing from l / 2 to 1 times as long as the seed proper. 
Distr. Malesia: Borneo (Sarawak), Philippines 
(Luzon), Celebes, Moluccas (Morotai, Halma- 
heira), New Guinea (incl. Misool and Louisiades: 
Rossel I.). Fig. 5. 




Fig. 5. Range of Crypteronia cumingii (Planch.) 
Planch, ex Endl. 



Ecol. Primary and secondary forests, on ridges, 
slopes, and along riverbanks, usually at low alti- 
tude, but also in the hills to 1200 m, once even 
found at 1800 m (Central Celebes, Malili area: 
B. Takale Kadju). 

Vern. Borneo: ubah, Iban, Sarawak; Philip- 
pines: andalai, Tag., ugdu, tigduon, Bik. ; Central 
Celebes, Malili area: kumba-a, langori tauru, 
longari, tomo wanna. 

Notes. As appears from his drawings, C. 
javanica Baill. must be referred here: bisexual 
flower with basal ovules ; very probably they were 
made after a Philippine specimen. 

In a depauperate collection from New Guinea 
(NGF 2958) the panicles are axillary. 

In specimens from the Moluccas frequently 
6-merous flowers occur among the normally 5- 
merous ones. 

In specimens from New Guinea there are usually 
some 4-celled ovaries and 4-celled fruits present 
among the predominantly 3-celled ones. 

3. Crypteronia griffithii Clarke in Hook. /. Fl. 
Br. Ind. 2 (1879) 574; King, J. As. Soc. Beng. 67, ii 
(1898) 5; Ridl. Agr. Bull. Str. & F. M. S. n.s. 1 
(1902) 180; Fl. Mai. Pen. 1 (1922) 821; Watson, 
Mai. For. Rec. 5 (1928) 176; Burk. Diet. 1 (1935) 
693; Corner, Ways. Trees (1940) 198. — Hens- 
lowia sp., Griff. Not. 4 (1845) 404; Ic. PI. As. 4 
(1845) t. 564, f. 1. — Fig. 6. 



1977] 



Crypteroniaceae (van Beusekom-Osinga) 



197 




Fig. 6. Crypteronia griffithii Clarkf , Singapore watcrcatchmcnt area, Dec. 1969 (Photogr. Van Beusekom). 



198 



Flora Malesiana 



[ser. I, vol. 8 2 



Tree up to 40 m, 60 cm 0. Twigs glabrous. 
Leaves elliptic-oblong to ovate-oblong, (5-) 10-25 
(-35) by 5-12(-15)cm, rounded or emarginate at 
the base, acute, sometimes acuminate at the top, 
tip usually obtuse, sometimes acute, glabrous, 
coriaceous, distinctly and slightly prominently 
reticulate beneath; nerves (8— )10— 1 5(— 18) pairs; 
petiole (2-)5-10mm, 2-5 mm 0, glabrous. 
Inflorescence axillary, one or a few together below 
the leaves on older nodes, 15-50 cm, only with 
primary side axes, very rarely with a few secondary 
side axes near the base, without distinct peduncle, 
sublignose, often ferrugineously pubescent, with 
pendulous axes; main axis terete or somewhat 
flattened, finely ribbed; primary axes 8-12(-15), 
not always paired, finely ribbed; racemules up to 
50 cm. Bracts of axes triangular, c. 1 mm, mostly 
caducous. Bracts of flowers subulate, c. 1 mm, 
persistent. Pedicels V2-K-IV2) mm, c. 1 / 2 mm 0. 
Receptacle c. 1 mm high, 2-3 mm wide, densely 
whitish papillose inside. Sepals triangular, 172-2 
mm. Stamens 5; filaments 3-4 mm, glabrous; 
connective V4-V2 mm, slightly conduplicated ; 
anthers linear, V4-V2 by c. 0.1 mm, ± latrorse. 
Ovary (sub)globose to pyramidal, 3-celled, c. 1 mm, 
more or less fine whitish papillose whether or not 
with longer hairs in between; style (lV2-)3-4(-5) 
mm, c. 1 U mm 0, sparsely puberulous; stigma 
slightly capitate. Ovules inserted on the somewhat 
conically elevated edges between the bases of the 
septa. Capsule (sub)globose, c. 2 mm; valves 3. 
Seeds 0.3-0.4 by c. 0.2 mm, apical and basal part 
of wing about IV2 times as long as the seed proper. 

Distr. Burma (Moulmein, one coll.); in Malesia: 
Central Sumatra (one coll.), Malay Peninsula 
(incl. Penang I.), Borneo (W. Sarawak, Sandakan, 
Gaya I., E. Kutai, Nunukan I.). Fig. 7. 

Ecol. In primary lowland forests, often on sandy 
soils, up to 500 m. 

Field notes. Bole usually straight. Bark surface 




smooth or rough and scaly. Twigs strongly swollen 
at the nodes. Young leaves purplish. Flowers dark 
blue to purple or magenta, sometimes noted yellow 
or yellowish reddish. 

Uses. Ridley (1902) reported the wood to be 
durable and used for house-building. 

Vern. Sumatra: panarahan; Malay Peninsula: 
bekwoi, kilat tampoi, nyirik bukit, simpo, sirumpu, 
simpoh, sumpu(t), telingga badak ; Borneo : engkolot, 
rambai rambai, Gaya I., ubah semut, I ban. 

4. Crypteronia macrophylla Beus.-Osinga, Blumea 
22(1975)261. — Fig. 1,2. 

Tree up to 20 (?) m. Twigs glabrous. Leaves 
ovate-oblong to lanceolate, (20-)25-45 by 7-15 cm, 
usually emarginate, sometimes rounded at the base, 
acute to ± acuminate at the top, tip obtuse, 
glabrous, coriaceous, distinctly and ± prominently 
reticulate beneath; nerves 15-25 pairs; petiole 
5-10(-15) mm, 2-5 mm 0, glabrous. Inflorescence 
ramifiorous, 45-90 cm, always branched up to the 
third order, without distinct peduncle, pendulous, 
woody, with spreading axes; main axis terete; 
primary axes many, not always paired ; racemules 
up to 30 cm. Bracts of axes triangular, c. 1 mm, 
caducous. Bracts of the flowers subulate, c. 1 mm, 
persistent. Pedicels 1-3 mm, c. 1 j 2 mm 0. Recep- 
tacle ± flat, l-P^mm wide, puberulous, not 
densely papillose inside. Sepals deltoid, c. 1 mm, 
inside with a longitudinal densely puberulous crest. 
Stamens 4; filaments 172-3 mm, glabrous, at the 
very base puberulous; connective c. 1 I 2 mm, more 
or less conduplicated; anthers ± linear, 72 by 
0.1-0.2 mm, ± introrse. Ovary pyramidal, 4-celled, 
1-2 mm, puberulous, more or less episeptally 
impressed; style (l-)l l /2— 2 1 / 2 (— 3) mm, c. l U mm 0, 
puberulous; stigma truncate, hardly wider than 
the style. Ovules inserted on the somewhat coni- 
cally elevated edges between the bases of the septs. 
Capsule (sub)globose to pyramidal, 2-272 by 
2-27 2 mm ; valves 4. Seeds c. l j 2 by c. i U mm, apical 
and basal part of wing as long as or slightly longer 
than the seed proper. 

Distr. Malesia: Borneo: Sarawak (Kuching, 
Sibu), Kutai (several localities), Sambas region. 
Fig. 8. 



^\~7^ 




Fig. 7. Range of Crypteronia griffithii Clarke. 



Fig. 8. Range of Crypteronia macrophylla Beus.- 
Osinga. 



Ecol. In primary forest up to 1200 m. Locally 
frequent. Young branches are often inhabited by 
ants. 



1977] 



Crypteroniaceae (van Beusekom-Osinga) 



199 



Field notes. Tree with semi-pendent branches. Note. The large complex, woody inflorescence 

Bark surface smooth. Young leaves rich purplish, which is always produced below the leaves on older 

Flowers greenish with purple-brown stamens, nodes is characteristic. 
Fruits dark green. 




Fig 9 Dacty locladu s stenoUachys Oli v. a. Habit, ' „ b. flower, ' 10, c. seed, 16 (a, b Fuchs 21 186, 

<S9261). 



2 DACTYLOCLAIM s 

Onv in HoolL Ic PI (1895) t. 2351; GlLG in E. & P. Nat. Pfl. lam. Nachtr. 1 
(1897) 267; Ham.../. Med. Rijkshcrb. 35 (1918) 18; Baku./. Rcc. Irav. Bot. 



200 



Flora Malesiana 



[ser. I, vol. 8 2 



Need. 40 (1943) preprint 329; Beus.-Osinga & Beus. Blumea 22 (1975) 261. — 
Fig. 9. 

Leaves elliptic or obovate to (obovate-)oblong, glabrous; midrib thickish, ± flat 
above, prominent beneath; nerves ± straight, anastomosing in an indistinct mar- 
ginal nerve. Inflorescence terminal or axillary to the highest leaves, erect, poorly 
branched ; axes flattened, puberulous ; racemules with at least a few tens of flowers. 
Flower-bracts caducous. Flowers bisexual, 5-(4-)isomerous, almost sessile. 
Receptacle in- and outside puberulous, accrescent. Sepals triangular, persistent. 
Petals unguiculate, with suborbicular, irregularly lobed lamina, in bud covering 
the stamens as a hood, soon caducous. Stamens persistent; filaments terete, some- 
what flattened; connective about orbicular, not conduplicate; anthers inserted 
transversally, somewhat below the upper margin of the connective, oblong to 
broad-linear, introrse. Ovary half-inferior, the lower part adnate to the recep- 
tacle, the top part semiglobose, puberulous, (3-), 4- or 5-carpellate, (3-), 4- or 
5-locular, septs not connate; style subulate, somewhat longer than the ovary, 
puberulous, persistent; stigma capitate. Ovules 3 per locule, inserted in vertical 
position, basally between the septs. Capsule almost inferior, broad-ellipsoid, small, 
pericarp chartaceous, puberulous, inside dehiscent down to the bottom with (3) 4 
or 5 valves of which only the upper 1 / 4 protrudes from that part of the pericarp that 
is surrounded by and fused with the enlarged receptacle, at the top often kept to- 
gether by the non-splitting stigma. Seeds 3 per locule (1 or 2 sometimes not 
developed), small, in vertical position; seed narrow-ellipsoid, flat, situated centrally 
in its more or less rectangular, narrow, membranous wing almost 2 times as long 
as the body of the seed; raphe running close to the embryo. 



Distr. Malesia: Borneo and W. New Guinea (sterile coll.). Fig. 10. 
Ecol. Lowland peat swamp forest. 



1. Dactylocladus stenostachys Oliv. in Hook. Ic. 
PI. IV, 4 (1895) t. 2351 ; Hall. /. Med. Rijksherb. 
35 (1918) 18; Merr. En. Born. (1921) 452; Diels & 
Hackenberg, Bot. Jahrb. 60 (1926) 312; Bakh./. 
Rec. Trav. Bot. Need. 40 (1943) preprint 329; 
Browne, For. Trees Sarawak & Brunei (1955) 261, 
t. 33; Anderson, Gard. Bull. Sing. 20 (1963) 178, 
pi. 1, 2, 6, 7; Meijer, Field Guide Trees W. Mai. 
(1974) 205, f. 51, pi. 14; Beus.-Osinga & Beus. 
Blumea 22 (1975) 262. — Fig. 9. 

Tree up to 40 m, dbh up to l'A* m, at the base 
producing pneumatophores. Twigs often several 
together per leaf-axil, the younger ones often with 
ribbed angles. Leaves 4— 8(— 16) by 2'/ 2 -4(-6) cm, 
with revolute margin, cuneate at the base, some- 
times emarginate, usually rounded up to acuminate 
at the top with acute tip, coriaceous; nerves 11-15 
pairs, usually rather obscure, flat to prominulent 
above and beneath; petiole 3-5 mm, 2-3 mm 0. 
Inflorescence when axillary 1-3 together, erect, up 
to 14cm, consisting of 3 racemules; peduncle up 
to 6cm, (sub)glabrous; axes finely ribbed; race- 
mules c. 8 cm, from 1 cm above the base ± densely 
set with flowers. Bracts of axes minute, soon cadu- 
cous. Bracts of the flowers linear or narrowly 
triangular, c. 1 mm. Pedicels up to V2 mrn - 
Receptacle c. 2 mm high, c. 2\' 2 mm wide. Sepals 
c. 1 mm. Petals c. 1 mm, puberulous outside and 



on the margin. Filaments c. 1 mm, 0.2 mm wide, 
puberulous; connective V4-V2 mm; anthers 0.2 by 
0.1mm. Style 172-2 mm, c . 0.2 mm 0; stigma 
V4-V2 nim. Capsule 3-4 by 2-3(-3V 2 ) mm - Seed c. 
1.4 by 0.2-0.3 mm, including wing 2V2-3 by 
"V.cT'At mm. 

Distr. Malesia: widely distributed in Borneo. 
It was by error incidentally reported from Malaya 
(PL Mai. Bull. p. 1696, p. 2375). Fig. 10. 

Ecol. Durant (For. Rep. Brunei, 1933, 6, 
photogr.) reported this species (under the name 
Crypteronia) to occur in Brunei as an associate of 










7^_ 



Fig. 10. Range of the genus Dactylocladus Oliv.; 
D. stenostachys Oliv. dots, D. sp. triangle. 



1977] 



Crypteroniaceae (van Beusekom-Osinga) 



201 



Dryobalanops and Combretocarpus in considerable 
quantity over large areas of freshwater swamp, 
often with 12 mature trees (over 30 cm 0) per ha. 
Diels & Hackenberg (I.e.) mentioned its occur- 
rence in the Sampit swamp forest area in SW. 
Borneo together with Combretocarpus, Campno- 
sperma, etc. Browne (I.e.) stated that it occurs in 
practically all types of peat swamp forest in Sara- 
wak. He found its frequency somewhat lower than 
Durant did; he found sometimes 8, but averagely 
3 mature trees per ha, but he said that locally, 
between Balingian and Bintulu, it was the dominant 
tree of the swamps of Sarawak. 

Browne recorded that the vernacular names 
jongkong and tabak allude to characteristic quali- 
ties Jongkong referring to the rather stout yellowish 
pneumatophores at the stem-base, while tabak 
would refer to the characteristic minute perfora- 
tions of the wood from radial vessels. 

Anderson (I.e.) confirmed Browne's observa- 
tion that it is one of the most characteristic swamp 
forest trees, the only species represented in all 
communities throughout Sarawak and Brunei. 
He produced photographs of the Gonystylus- 
Dactylocladus-Neoscortechinia (I.e. pi. 1 & 2) and 
the Combretocarpus-Dactylocladus associations 
(I.e. pi. 6 & 7). 

Its wide range in the peat swamps is also con- 
firmed palynologically in the peat according to 
Anderson & Muller (Rev. Palaeobot. & Palyn. 
19, 1975, 314-316) where it figures in phases 1-6. 
In a Miocene deposit in Brunei, near Berakas, 
Muller reported also its occurrence (I.e. f. 5, 
diagram) with the associates as today. It must be 
added that its pollen can be confused with that of 
Axinandra and some Melastomataceae of which 
the latter also occur in peat swamps, albeit in small 
numbers. 

It is remarkable that whereas this type of peat 
forest ranged unbrokenly at least from the 
Miocene to the Present, Dactylocladus is not yet 
found in Sumatra and Malaya, provinces with 
which Borneo was joined, during the Pleistocene 
Glacial period, by a huge lowland riverine area 
(now the South China Sea); this would have made 
dispersal and exchange very probable, as it was for 
its associate Combretocarpus and some species of 



Gonystylus. It may yet be discovered in the peat 
swamps of Malaya and Sumatra, but must then be 
very rare. 

Uses. According to Browne (I.e.) it is the fourth 
important export timber tree of Sarawak. Extrac- 
tion is facilitated by the fact that the logs float in 
water. Meijer (I.e.) termed it a general utility 
timber in Sabah where it is, besides Gonystylus, 
the most important export timber from the Klias 
Peninsula. 

Vern. Jongkong, tabak (the most common 
names), entibu, garu buaja, jinjang, (medang) 
b&ladi, m. miang, melinkat kerangas, miribong, 
tirenjangan. 

Dactylocladus sp. — Cf. Meijer, Field Guide Trees 
W. Mai. (1974) 205, in note. 

Distr. Malesia: West New Guinea: in peat 
forest along Rouffaer R., the only large tree in this 
forest type, c. 175 m alt., Docters van Leeuwen 
9973 (BO, L, etc.), sterile, distributed as Memecylon 
sp. Fig. 10. 

Notes. Through Meijer (I.e.) attention was 
drawn to this collection which was pre-identified 
through the uncanny form knowledge of Mr. Nedi 
at Bogor. Mr. G. van Vliet, Leyden, has examined 
the leaves anatomically and found no difference 
with the Bornean species. Dr. van Steenis, who 
unearthed the Leyden duplicate, has found that it 
shares a small but significant vegetative character 
with the Bornean species, viz the occurrence of a 
shallow, rimmed cavity-like depression at the 
extreme base of the petiole, similar to that in e.g. 
Garcinia; the two small cups together envelop the 
terminal bud. This is not found in Memecylon. 

Though in fact he found the sterile macromor- 
phology exactly matching, we like to postpone 
judgement on specific status until flowers and fruit 
are available. 

About the considerable geographical gap be- 
tween Borneo and West New Guinea it can be said 
that true peat forest is not known from this gap 
at the present time. This is no explanation, how- 
ever, as the same gap occurs in the genus Koom- 
passia (Leg.-Caes.) which is not a peat-forest tree 
genus; in that genus the New Guinean species is 
distinct from the two of the Sunda shelf. 



3. AXINANDRA 

Thw. in Hook. J. Bot. 6 (1854) 66, t. 1 C; En. PI. Zeyl. (1859) 122; B. & H. Gen. 
PI. 1 (1867) 784; Bedd. Fl. Sylv. 2 (1869) t. 207; Baill. Adansonia 12 (1876) 84; 
Clarke in Hook./. Fl. Br. Ind. 2 (1879) 581; Cogniaux in DC. Mon. Phan. 7 
(1891) 1113;KRASSERin E. & P. Nat. Pfl. Fam. 3, 7 (1893) 142, 196; Bakh./. Rec. 
Trav. Bot. Neerl. 40 (1943) preprint 332; Meijer, Ceyl. J. Sc. (Biol. Sc.) 10 (1972) 
72; Beus.-Osinga & Beus. Blumca 22 (1975) 262. — Naxiandra (Baill.) Krasser 
in E. &P. Nat. Pfl. lam. 3, 7 ( 1 K93) 197, f. 182 A. — Fig. 11. 

Leaves elliptic to oblong, sometimes ovate, glabrous; midrib impressed above, 
prominent beneath; nerves ascending and anastomosing in a looped marginal 
nerve, intramarginal nerve present. Inflorescence terminal or axillary and then at the 
end of the twigs, erect, poorly branched; axes more or less angular, puberulous; 
racemules with up to some tens of flowers. I lower-bracts 3 per flower, the outer 



202 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 11. Axinandra coriacea Baill. a. Habit, x 1 I 2 , b. venation undersurface of leaf, nat. size, c. bud, 
d. older bud, style protruding, petals separating, e. mature flower, petals dropped, f. old flower, petals and 
stamens dropped, all x l l j 2 , g. ripe capsule, x 3 / 4 , h. stamen, x 15 (a SAN 57276, c-f, h S 14489, g 

Meuer SAN 49845). 

ones often minute, persistent during anthesis. Flowers bisexual, 5(-4)-merous, 
pedicelled. Receptacle puberulous outside only, glabrescent in fruit, much accres- 
cent and lignified in fruit. Sepals 5 (4), deltoid, more or less acuminate, evanescent 
in fruit. Petals 5 (4), valvate-connate or valvate- (or somewhat imbricate-)condupli- 
cate or imbricate-contorted, more or less connate or cohering and soon dropping 
simultaneously in the shape of an umbrella, flimsy, consisting of a wide basal part 
tapering into a narrower median part which widens into a whether or not well- 
developed, frayed apical part, induplicate and enveloping the stamens which are 
situated in pairs between two longitudinal lamellae on the inside of each petal. 
Stamens 10 (8), epi- and alternisepalous, (sub)equal, caducous; filaments (rather) 
thick and short, terete, to more or less flattened, tapering upwards; connective 



1977] Crypteroniaceae (van Beusekom-Osinga) 203 

wide, large, conduplicate, mostly provided with a more or less distinct dorsal 
tubercle; anthers marginally at the apex of the connective, broad-linear, introrse. 
Ovary inferior, immersed in the receptacle, 3-(2-)carpellate, 6-(4-)celled, with 
free or towards the base somewhat connate septs, glabrous; style (subulate-) 
terete, shorter to distinctly longer than the receptacle, and whether or not pro- 
truding from the ripe bud, glabrous, evanescent in fruit; stigma punctate, minute. 
Ovules 1 or 2 per cell, two (one) by two (one) in vertical position inserted basally 
on either side of the 3 (2) stouter ones of the 6 (4) septs. Capsule half-inferior, big 
and woody, globose to ellipsoid, (sub)glabrous, when young provided with a frag- 
ment of the style, inside dehiscent down to the bottom with 2-6 valves of which 
only the upper parts protrude from that part of the capsule which is surrounded by 
and fused with the enlarged receptacle of which the rim often remains visible as an 
irregular more or less conspicuous rib; visible part of the valves triangular, coarse, 
solid, deltoid at cross-section, glabrous. Seeds few, in vertical position; seed 
depressed-ellipsoid, situated basally and obliquely in its thin, (narrow-)oblong 
wing 2-3 times as long as the body of the seed; raphe running from the basal inser- 
tion all along the wing margin back to the embryo. 

Distr. Ceylon (1 sp.) and Malesia: Malay Peninsula (one record) and Borneo (3 spp.) but nowhere 
common. Fig. 12. 

A. zeylanica was also mentioned to occur in Borneo by Bakhuizen/. (Rec. Trav. Bot. Need. 40, 1943, 
preprint 332) but Hallier/. 2683, on which this was based, belongs to A. coriacea. The same author (I.e.) 
recorded A. borneensis Bakh./. (= A. beccariana) from Billiton I. ; the sheet Teysmann s.n. was collected, 
however, on Mt Blitong in Borneo. 

As with Dactylocladus the extreme rarity of the genus on the Sunda-shelf west of Borneo is remarkable 
and remains unexplained; sofar known Axinandra is not bound to a rare or very specialized biotope. 

Ecol. Lowland and submontane rain-forest, up to 1200 m. 

KEY TO THE SPECIES 

1 . Connective pear- or drop-shaped. Filaments 1 l l 2 -2 mm. Style 2-5 mm. Ovules 2 per cell. Sect. Naxian- 

dra Baill. 
2. Internodes winged upwards. Leaf-base rounded to emarginate. Inflorescences distinctly longer than 

5 cm 1. A. alata 

2. Internodes not winged. Leaf-base shortly attenuate. Inflorescences shorter, or distinctly longer than 
5 cm. 
3. Leaves chartaceous. Tip of leaf-apex acute. Inflorescence 5-20 cm. Petals c. 3 mm. Fruit IV2-2 by 

1-1 ' 2 cm; rim of enlarged receptacle about median 2. A. beccariana 

3. Leaves coriaceous. Tip of leaf-apex obtuse. Inflorescence stunted, (l-)2-4(-5) cm. Petals c. 6 mm 
(fig. lid). Fruit 2-3(-3'/2) by \ l l 2 -2 l l 2 cm; rim of enlarged receptacle infra-median (fig. llg) 

3. A. coriacea 

1. Connective quadrate-elliptic. Filaments c. V2 mm. Style shorter than '/a rnrn- Ovules 1 per cell. Sect. 

Axinandra. Species of Ceylon A. zeylanica Thw. 

1. Axinandra alata Baill. Adansonia 12(1876) 86; bearing 10-35 flowers. Bracts of axes deltoid to 

Bull. Soc. Linn. Paris (1877) 128; Cogniaux in triangular, c. I mm, acute at the top, minutely 

IX Mon. Phan. 7 (1891) 1114; Mirk. En. Born, pubcrulous, mostly caducous. Floral bracts small, 

(1921) 452; Bils.-Osinga & Beus. Blumea 22 the middle one only slightly exceeding the lateral 

(1975) 263. ones, narrowly triangular, c. I mm. Pedicels 

Tree. Twigs glabrous; internodes terete at the 1-2 mm, c. '/2 mm 0. Receptacle c. 2 mm high, 

base, growing acutely quadrangular upwards with c. 3 mm wide, ribbed, densely puberulous. Sepali 

4 gradually widening wings towards the nodes, ft ' , mm. Petals C. V , 4 mm, I I 1 4 mm wide at 

(suh)glabrous; wings I 3 mm (<>r nunc') wide at the base, valvatc to imbricate at the base, valvate- 

the top, ending in an acute processus often curved conduplicate for the rest, more or less connate 

upwards, wearing off when older, /caves J 10 hy especially in the median part, almost completely 

3 5 cm, rounded at the base, cuspidate at the top enveloping the stamens. Itlamcnts c 2 Rim, 

and with acute tip, chartaceous to subcoriaceous, ' 4 mm wide at the base, glabrous. ( omu 

distinctly reticulate; nerves c. 12 pans, Hat above, pear- or drop-shaped, r. I mm, with dorsal swelling, 

prominent beneath ; petiole 1 5 mm, I I "j mm 0. Anther-cells c. I mm, c. ', mm wide. Style 3 5 Rim, 

hrflorescene* % 13 cm, consisting of mam axis with '/« mm 0, DTOtniding from mature bud. Ovules 

I or 2 pairs of 4 12 cm long primary axes each 2 per cell. Iruit not seen. 



204 



Flora Malesiana 



[ser. I, vol. 8 2 



Distr. Malesia: Borneo (Sarawak), only 
known from the type. 




A. maingayi). The differences between this speci- 
men and the material from Borneo as mentioned 
by Clarke I.e. are in my opinion of minor impor- 
tance; consequently, I have included A. maingayi 
in the synonymy of A. beccariana. 



Fig. 12. Range of the genus Axinandra Thw. In 

Ceylon 1 sp., in Borneo 3 spp. of which one once 

found in Malaya. 

2. Axinandra beccariana Baill. Adansonia 12 
(1876) 85; Bull. Soc. Linn. Paris (1877) 127; 
Cogniaux in DC. Mon. Phan. 7 (1891) 1114; 
Merr. En. Born. (1921) 452; Beus.-Osinga & 
Beus. Blumea 22 (1975) 263. — A. maingayi 
Clarke, Fl. Br. Ind. 2 (1879) 581 ; Cogniaux in 
DC. Mon. Phan. 7 (1891) 115; Ridl. Fl. Mai. Pen. 
1 (1922) 826. — A. borneensis Bakh. /. Rec. Trav. 
Bot. Neerl. 40 (1943) preprint 332. 

Tree. Twigs glabrous or very minutely puberu- 
lous. Leaves 5-10 by 2-5 cm, shortly attenuate at 
the base, acuminate to cuspidate at the top, with 
acute tip, chartaceous, faintly reticulate; nerves 
8-12 pairs, fiat above, prominent beneath; petiole 
5-8 mm, 1-2 mm 0. Inflorescence 5-20 cm, 
branched up to the second order, with slender axes 
each bearing up to c. 1 5 flowers ; primary axes up to 
4 pairs, 4-15 cm. Bracts of the axes deltoid to 
narrowly triangular, 1-3 mm, acute at the top, 
subglabrous, mostly caducous, sometimes partly 
present as small or reduced leaves. Floral bracts 
small, the middle one triangular to linear-lanceo- 
late, 7 2 -4 mm, the lateral ones minute. Pedicels 
1-2V 2 rnm, c. 7 2 mm 0. Receptacle c. 2 mm high, 
c. 3 mm wide, sometimes faintly ribbed, sparsely 
minutely puberulous. Sepals c. 1 t 2 mm. Petals c. 
3 mm, c. 1 mm wide at the base, valvate, almost 
completely connate, only enveloping the dorsal and 
part of the apical side of the stamens. Filaments c. 
P^mm, c. V2 mm w 'de at the base, glabrous. 
Connective pear- or drop-shaped, c. F/2 mm , with 
dorsal swelling. Anther-cells c. 1 by c. 1 I 4 mm. 
Style c. 2 mm, c. l j 2 mm 0, hardly or not protrud- 
ing from mature bud. Ovules 2 per cell. Capsule 
172-2 by l-17 4 cm, faintly 10-ribbed, rim of 
enlarged receptacle about median ; valves c. 3 / 4 cm. 
Seeds not seen. 

Distr. Malesia: Borneo and Malay Peninsula 
(Malacca, one old record). Fig. 13. 

Ecol. Lowland forests. 

Note. From the Malay Peninsula only one 
collection is known (Maingay 654-2, type of 




Fig. 13. Localities of Axinandra beccariana Baill. 
(triangles) and A. coriacea Baill. (dots). 

3. Axinandra coriacea Baill. Adansonia 12 (1876) 
85; Bull. Soc. Linn. Paris (1877) 127; Hist. PI. 7 
(1880) 28, f. 43; Cogniaux in DC. Mon. Phan. 7 
(1891) 1114; Merr. En. Born. (1921) 452; Beus.- 
Osinga & Beus. Blumea 22 (1975) 264. — Naxian- 
dra coriacea (Baill.) Krasslr in E. & P. Nat. Pfl. 
Fam. 3, 7 (1893) 197. — Fig. 11. 

Tree up to 35 m, 50 cm 0. Twigs glabrous. 
Leaves 5-10(-12) by (17 2 -)2-5(-6) cm, shortly 
attenuate, sometimes acute at the base, acuminate 
to cuspidate at the top, with obtuse tip, coriaceous, 
conspicuously reticulate; nerves 8—12 pairs, fiat 
above, prominent beneath ; petiole 5-8 mm, c. 
1-2 mm 0. Inflorescence ( 1 -)2— 4(— 5) cm, con- 
sisting of a few, sometimes one 1-5 cm long rather 
coarse, sometimes stunted axes, each bearing 0-15 
flowers. Bracts of the axes deltoid to triangular, 
c. 1 mm, obtuse at the top, subglabrous, mostly 
caducous. Floral bracts small, the middle one del- 
toid to triangular, c. 1 mm, the lateral ones minute. 
Pedicels 1— 2 1 / 2 mm, c. 1 mm 0. Receptacle c. 3 mm 
high, 3-4(-5) mm wide, densely puberulous. 
Sepals c. 1 mm. Petals c. 6 mm, c. 17 2 mm wide at 
the base, valvate at the base, conduplicate-valvate 
for the rest, more or less connate in the median 
part, mostly almost completely enveloping the 
stamens. Filaments \ l \ 2 -2 mm long, 3 / 4 -l mm wide 
at the base, glabrous. Connective pear- or drop- 
shaped, c. 17 2 mm, with dorsal swelling. Anther- 
cells c. 17 2 rnm, c. 74 mm wide. Style 2-5 mm, 
c. 7 2 mm 0, protruding from mature bud. Ovules 
2 per locule. Capsule 2-37 2 by 172-272 cm; rim of 
enlarged receptacle infra-median ; valves 1 / 2 — 1(— 1 72) 
cm. Seeds 0.7-0.8(-l) by 0.3-0.4(-0.5) cm; wing 
c. l l 2 cm wide. 

Distr. Malesia: Borneo. Fairly rare. Fig. 13. 

Ecol. Primary (dipterocarp) forest at low and 
medium altitude, up to 1200 m, also recorded 
from ultrabasic red-brown soil. 

Field notes. Buttresses when present up to 1 l j 2 - 
2 m vertically. Bark surface flaky. Stamens pale 
yellow. Flowers greenish; corolla white. 

Vern. Ubah, Iban, Sarawak, obah, Sabah. 



SYMPLOCACEAE (H. P. Nooteboom, Leyden) 1 

The family consists of one genus only, Symplocos, which occurred already in the 
Eocene over the entire northern hemisphere in the mixed mesophytic forest and in 
all probability also in the Indo-Australian tropics. 

As proved by abundant fossil endocarps, the Eocene species had already a fruit 
structure very similar to that of now living species and the genus existed at that early 
time obviously already in optima forma, a reason to assume that it must be of high 
antiquity. This is also corroborated by the fact that the tropical subgenus Symplo- 
cos has a very disjunct trans-Pacific range; explanation by chance transoceanic 
long-distance dispersal must be refuted because it is in contradiction with all 
presently known facts. 

Although Symplocos has shown a fairly abundant speciation, considering its 
present size and 25 fossil species described, it has surprisingly not led to other 
generic development and remained in splendid isolation. 

Its systematic affinities induced mostly to classify it with Ebenales. In my mono- 
graph of the Old World species (1975) I have brought all evidence together and 
have concluded that this position is unlikely: pollen structure differs from that in 
other families of Ebenales, so do the stomata, the placentation and the structure of 
the ovules. This leads to the view that Symplocos is more allied to Comaceae and 
Theaceae, sharing also with both families a primitive wood anatomy. Still the 
affinity is not that close, as for example Theaceae have a truly axile placentation. 
The chromosome number fits better with Comaceae sens. lat. 



SYMPLOCOS 

Jacq. En. Fl. Carib. (1760) 5, 24; Select. Stirp. Am. Hist. (1763) 166, t. 175, f. 68; 
Linne, Gen. PI. ed. 6 (1764) 272; Miers, J. Linn. Soc. Bot. 17 (1879) 285; Brand, 
Pfl. R. Heft 6 (1901) 13, 9 fig.; Noot. Leid. Bot. Ser. 1 (1975) 33, 7 fig., 21 pi., with 
full synonymy. — Fig. 1-20. 

For synonyms see under the subgenera. 

Shrubs to (rarely) large, (in Mai.) evergreen trees; bark in various spp. bitter; 
growth continuous or interrupted (in flushes), in the latter case the buds protected 
by often leathery bud-scales; glabrous or hairy (by simple hairs). Leaves simple, 
alternate or spirally arranged, rarely pseudoverticillate, estipulate, penninerved, 
petioled, rarely almost sessile ; when dry often discolouring (often in yellow tinges)in 
subg. Hopea. Flowers in spikes, racemes, or panicles, mostly from the upper leaf- 
axils, sometimes condensed to clusters, sometimes terminal or from the axils 
of fallen leaves, rarely solitary; supported by a bract and 2 bracteoles, rarely 
several bracts and bracteoles by abortion of flowers; flowers actinomorphic, 
bisexual, rarely by reduction unisexual and plant polygamous, not rarely 
fragrant, distinctly so in subg. Symplocos. Calyx with a very short tube above 
the inferior ovary, the limb 3 5-lobed, imbricate, persistent, sometimes split into 
two parts and seemingly 2-lobed. Corolla sympetalous, but divided nearly to the 
base in subg, Hopea\ lobes (3 )5( 10 in the New World), c|uineuneially imbricate, 
whitish, bluish or purplish. Stamens 4 to mostly x, eonnate in a long monadel- 

(i) With co-operation <>i the General i ditor. 

(205) 



206 



Flora Malesiana 



[ser. I, vol. 8 2 



phous tube, at its base adnate to the corolla and very unequal, but in subg. Hopea 
only connate at the very base, monadelphous or pentadelphous and then the 
bundles alternipetalous; anthers globose, 2-celled, lengthwise dehiscent, introrse. 
Ovary inferior (to ± semi-inferior), 2-5-celled, with a complete septation; style 1, 
stigma punctiform or peltate. Ovules 2-4 in each cell, pendulous, anatropous- 
epitropous or amphitropous, unitegmic, tenuicellular. Drupe monopyrenous, 
crowned by the persistent calyx lobes, of various shape: cylindrical to globose, 
ampulliform or spindle-shaped; mesocarp usually thin, sometimes thick and then 
often quite hard; stone smooth or mostly sculptured in various degree or length- 
wise ridged. Seeds straight or curved, 1 in each developed cell, with copious 
endosperm ; embryo straight or curved, with very short linear cotyledons. 

Distribution. About 250 spp. , in the eastern parts of the Old World, from Ceylon and Bom- 
bay in the Deccan to Fiji in West Polynesia and from Manchuria at 46° N as far as New South 
Wales and Lord Howe I. at 32° S; in the New World from the State of Washington in the U.S.A. 
to S. Brasil; throughout Malesia. Fig. 1. 




a Pliocene 

o Miocene 

• Oligocene 

+ Eocene 



Symplocos subg. Hopea 'ti 

Symplocos subg. Symplocos 



Fig. 1. Range of the genus Symplocos, recent and fossil. The fossil localities in Europe, Japan, and E. 
North America are all belonging to species of subg. Hopea. 



There is no species common to the Old and New World, but the E. Asian S. lucida is closest 
allied to the N. American S. tinctoria. 

Taxonomy. Brand (1901) has made an intricate subdivision of the genus, partly based on 
former generic names. I believe we cannot go further than a subdivision into two subgenera, in 
which macromorphology is supported by chemotaxonomy and palynology, viz subg. Symplocos 
and subg. Hopea. 

Subdivisions could be based on one important single character: straight versus curved embryo, 
spiral versus distichous phyllotaxis, continuous versus flushwise growth from scaly buds, but it 
appears that such subdivisions do not coincide. This leads to the view that there is a block of 
species with reticulate affinities. This view also emerges from the palynological results. 



1977] Symplocaceae (Nooteboom) 207 

Both subgenera occur in the New and the Old World ; subg. Symplocos, which is almost strictly 
tropical, possesses only 2 spp. in Indo-Malesia, but probably many more in America. 

In this revision 58 spp. are distinguished in Malesia; there are more new species, but I have 
refrained from describing them as the material is incomplete; I have enumerated them in my 
revision Ac. 296. 

Fossils. Before the Glacial Epoch Symplocos occurred also in Europe in the mixed mesophytic 
subtropical to warm-temperate forest, onwards of the Eocene, obviously as a common con- 
stituent of the Tertiary mixed mesophytic forest, as shown from fossil stones. Cf Kirchheimer, 
Palaeontographica 90B (1949) 1-52, t. 1-2. These stones are very similar to endocarps of recent 
species; obviously no major changes did occur in the genus during this era. The three fossil 
Pliocene species in Japan are almost certainly the same as those that are living there today. One 
fossil species is known from the Eocene in the eastern U.S.A. Fig. 1. 

Ecology. All species are evergreen, except a single deciduous one, S. paniculata (Thunb.) 
Miq. from Kashmir to Manchuria and Japan. 

They grow under tropical to temperate conditions in mixed evergreen rain-forest, not under 
arid conditions. 

Their stature is mostly small and they make part of the undergrowth and lower storeys, in 
exceptional cases attaining a maximum height of c. 30 m and 60 cm 0. 

In Malesia they are found from sea-level up to the alpine zone at c. 4000 m (Mt Kinabalu ; New 
Guinea), where they are represented by mostly microphyllous (fig. 12) dwarf shrubs in the dense 
elfin and mossy forest on slopes, summits and ridges where they may be common ; but they are 
almost nowhere recorded as a dominant. 

A few species, e.g. S. polyandra, are restricted to the lowland, but most species have a fair 
altitudinal range, and are most commonly collected in the hill and mountain forest. A few are 
restricted to high altitude, e.g. S. buxifolia, S. deflexa, S. johniana, S. zizyphoides, and several 
varieties of S. cochinchinensis. 

A fair number seem to be rare and have been seldom collected, others are common and widely 
distributed in the archipelago, notably S. cochinchinensis, S. celastrifolia, S. fasciculata, S. 
laeteviridis, S. ophirensis, and S. odoratissima. 

Especially these species, several of which are variable, grow on a variety of soils, including 
young-volcanic; they are scarce on limestone and generally prefer more acid, humous soils, e.g. 
S. celastrifolia is common in coastal forests, especially in the transition between mangroves and 
freshwater swamps, but it occurs also on kerangas, along river banks, and even in peat swamp 
forest. 

S. cochinchinensis var. sessifolia is very resistant against poisonous crater gases and acid soil 
conditions and can act as a pioneer in crater fields in Java, sometimes dwarfing down to very 
small size, although still producing flower and fruit; in the surrounding closed elfin forest it is a 
common small tree, growing together with Vaccinium, Myriea, Myrsine, Leptospermum, etc. 

Density of species. In fig. 2 the density of species has been indicated for each province and island 
(group). The richest areas are those of continental SE. Asia and West Malesia, while the number 
of species tapers out towards East Malesia and the SW. Pacific. The greatest number of endemic 
species is found in West Malesia, notably (as usually) in Borneo and the Philippines. However, in 
East Malesia New Guinea has a fair number of endemic species. The high number of endemics in 
New Caledonia is a bit exaggerating the situation as all are certainly derivatives of S. cochin- 
chinensis. The same holds for the endemics of New Guinea (with the exception of S. cylindracea) 
and for Australia (with the exception of S. cyanocarpa C. T. White). 

Flower biology. In all Symplocos spp. the flowers of an inflorescence open almost simultaneously 
and on one tree almost all inflorescences are open at the same time, so that the whole crown is 
for a short time gay with the blossoms (fig. 3). Of S. cochinchinensis var. sessifolia flowers are 
deliciously scented, as hawthorn, but field records mention other species as scentless or faintly 
scented. This varies obviously with the species. 

Pollination. Docters van Leeuwen (Verh. Kon. Ak. Wet. A'dam sect. 2, 31, 1933, 218) 
reported of .V. cochinchinensis var. sessifolia, on the summit of Mt Pangrango, West Java, at 
c. 3000 m, that flowers expand in the morning but open only halfway, the corolla remaining bent 
over the sexual organs; at 8 h. anthers are open and often touch the stigma on which the sticky 



208 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 2. Density of species in Old World Symplocos; above the hyphen the endemic species for each island 
(group) or country, below the hyphen the non-endemic species. 




Fig. 3. Symplocos laeteviridis Stapf var. laeteviridis in full flower, showing also alternate phyllotaxis.- 
Sabah (Nooteboom 1017). Photogr. Nooteboom, Febr. 1969. 



1977] Symplocaceae (Nooteboom) 209 

pollen readily falls; on the 2nd flowering day the corolla is widely open, anthers are empty, and 
the stigma is always pollinated. This means self-pollination. Docters van Leeuwen found, 
however, also the flowers frequented by various insects, among them bees and bumble-bees. They 
are not so much attracted by the little nectar, but are in search of pollen. 

Some species may have locally a strict flowering time; e.g. S. cochinchinensis var. sessifolia 
flowers, according to Docters van Leeuwen (I.e., fig. 52), from October to January, in the rainy 
season, on the summit of Mt Pangrango, West Java. 

Hybridization. Though there are in a few instances indications (by high sterile pollen %) that 
hybridization may occur, no clear cases are recorded. It is, however, clear that cross-fertilization 
must occur in the polygamous species in East Malesia. 

Galls. Docters van Leeuwen (Zoocecidia, 1926, 460) found in S. cochinchinensis var. sessi- 
folia small leaf galls, caused by psyllids by which the two halves of the leaf curve upwards till 
margins touch and a narrow cavity is formed. He recorded similar galls also from other forms of 
this species. In S. fasciculata he found a stem gall caused by a gall-midge and in S. brandisii a 
flower gall caused by a gall-midge. 

Dispersal. Ridley (Disp. 1930) assumed that bats may be fond of the hard-fleshed drupes 
(I.e. 347). He mentioned that in North America tyrant birds (Sayornis phoebe) eat amongst others 
fruit of S. tinctoria {I.e. 483) and that in South America a curassow, a sort of turkey, would feed 
on the fruit of 5. cernua. Docters van Leeuwen (Verh. Kon. Ak. Wet. A'dam sect. 2, 31, 1933, 
220) believed Symplocos to be dispersed by birds but did not find endocarps in the stomach of 
fruit-eating birds. Van Steenis found fruit of S. henschelii abundant on the ground below trees at 
Tjibodas, although this species has a fairly thick, hard-fleshed exocarp, in contrast to most 
species in which the exocarp is thin. Also in fossils sometimes immense quantities of stones are 
found together, about which Kirchheimer reported (Palaeontographica 90B, 1949, 1-52): in a 
total mass of c. 3500 m 3 he estimated the number of endocarps at some 2 l l 2 billions. He assumed 
that these were deposited within one century in a site of forest dominated by Symplocos. However, 
he added that the layers in which the endocarps were deposited gave no evidence of rivers which 
could have transported and accumulated the seeds and he concluded that they have dropped to 
the soil in situ. For these reasons abundant dispersal by birds or bats is in Symplocos not very 
likely. 

Dispersal by water takes place in species in which some fruit cells are barren and remain empty, 
e.g. S. celastrifolia. 

Morphology. The phyllotaxis is variable but constant for the species; it is either spiral or 
alternate (distichous) in which latter case the twigs are often zigzag (fig. 3). 

In most species leaves are more or less equally dispersed along the twigs, but in other species 
there is a tendency that the leaves are becoming crowded towards the end of the year's growth, 
e.g. in S. macrocarpa, as noted by Trimen (Handb. Fl. Ceyl. 3, 1895, 103). In Malesian spp. this 
occurs also in S. herzogii and S. gigantifolia where the large leaves occur crowded at the end of the 
year's growth. 

There is a single species in which all the leaves are in real pseudo-whorls, viz S. verticillifolia 
from the Philippine Is. (fig. 20). 

The leader-shoots in Symplocos, e.g. S. fasciculata, have spiral phyllotaxis; such shoots may, 
however, also carry flowers. 

Rejuvenation is in certain species by continuous growth of the twig apex, as is e.g. charac- 
teristic in S. fasciculata. In other species, however, there are clear buds with conspicuous bud- 
scales, indicating that the growth mode is flushwise and discontinuous, as e.g. in S. costata and 
S. lucida (fig. 15). This might be a good character of subdividing subg. Hopea. It can, however, 
only be used if one has accurate knowledge of the rejuvenation process of each species. This is 
sometimes difficult to ascertain from herbarium material as the bud-scales do not always leave 
traces of distinct scars, field data hardly ever mention the character, and material is seldom 
collected in the stage of flush. If the growth mode were well examined in all species I believe it 
would represent a good key character. 

f lushwise, discontinuous growth, with scaly buds could be assumed to be an adaptation to 

seasonally cold climates, it is a life form intermediate halfway evergreen ami deciduous, it is rare 

in the Malesian tropics v. here it is known e.g. from Acer, some genera of Lauraceae, Fagaceae, 



210 



Flora Malesiana 



[ser. I, vol. 8 2 



which also in the tropics are found in the cool, tropical-montane climate, which is however hardly 
seasonal. It still could be viewed as an indication of former immigration of taxa of higher latitude. 
Once acquired this growth mode must then have been conserved, as it occurs also in S. barring- 
toniifolia which is restricted to the tropical lowland. 

The inflorescence is either a panicle or a raceme or spike. Morphologically it is cymose, the 
flower always being sustained by two bracteoles which may at times carry abortive buds in their 
axil (fig. lib). In some cases the inflorescence is condensed to a fascicle or cluster of flowers 
(fig. 20a) or even be reduced to a single flower (fig. 19a). In a few species flowers occur on old 
wood, as e.g. in S. polyandra, S. wikstroemifolia (p.p.), S. rubiginosa, and S. tricoccata. 

The flowers are bisexual but functionally unisexual flowers are found in several taxa, especially 
in New Guinea. Such taxa are either dioecious or polygamous. In male flowers the style is small 
and without a stigma, in female flowers the number of stamens is reduced (even to less than 10) 
and anthers are sterile. In subg. Symplocos the stamens are monadelphous with a long tube 
(fig. 6a, d); in subg. Hopea they are only connate at the base for at most 2 mm (fig. lie), and 
intergrading from strictly monadelphous to strictly pentadelphous, the phalanges being alterni- 
petalous. 

In my revision it has been explained that, in contrast with former opinion, the ovary is initially 
1 -celled, with the ovules attached close to the centre on the induplicate part of the carpels, each 
of the 2-5 compartments having usually 4 ovules; in fruit these appear as cells. In each developed 
cell there is usually one seed. The latter and the embryo it contains may be curved or straight. 
See fig. 4b, c, g, h, j, k. 





Fig. 4. Symplocos ophirensis Clarke ssp. perakensis (K. & G.) Noot. var. perakensis. a. LS of fruit, out of 
centre, b. seed, c. curved embryo, with 2 short apical cotyledons, all x 4. — S. ophirensis Clarke ssp. 
cumingiana (Brand) Noot. var. cumingiana. d. LS of fruit, seed cavity empty, e. stone, x 4. — S. macro- 
phylla Wall, ex DC. ssp. cordifolia (Thw.) Noot. var. apicalis (Thw.) Noot. f. Ribbed stone, with fold, 
g. seed, the curved embryo enveloped by the albumen, x 2. — S. paniculata (Thunb.) Miq. h. LS of seed 
showing curved embryo, h'. ditto in CS, showing how such seed may appear deceptively as 2 seeds, x \ l j 2 . 
— S. glauca (Thunb.) Koidz. i. Fruit, j. seed in LS showing straight embryo, x 3. — S. paniculata 
(Thunb.) Miq. k. Germinating seedling with LS of endocarp and seed, showing mode of exist of embryo, 
x 1V 3 , 1. seedling, x 2 / 3 . — S. celastrifolia Griff, ex Clarke, m. U-shaped seed, x 6 (a-c Burkill 1013, 
d-e Nooteboom 2229, f-g Ashton 2480, k-l after Lubbock). — alb albumen, ec stony endocarp, mc 
mesocarp, os outer surface of fruit, sc seed cavity. 



1977] Symplocaceae (Nooteboom) 211 

The fruit is a drupe, with a fleshy, corky or woody mesocarp and a very hard stone (endocarp). 
The endocarp may be smooth (fig. 10c, 19d) or show outside ridges or irregularities (fig. 4e, f, 9c, 
lOe, 14d); the same holds for the inside of the endocarp. In the centre of the copious endosperm 
the embryo is embedded. It is slender and may be straight or curved. In the tropical subg. 
Symplocos it is always straight. In subg. Hopea it is straight in all American spp. and in 80% of 
the living species in the Old World and also in all fossil species in Europe. From this it is con- 
cluded that a straight embryo seems to be the primitive state in the genus. Only the three Pliocene 
fossil species of Japan, which can be matched with living species, have curved seeds and conse- 
quently curved embryos. 

Curved seeds occur in degree, they may be hook-shaped or U-shaped or even be twice curved 
(S-shaped in S. brachybotrys). See fig. 4. This may give some difficulty in studying sections of the 
stones to count the number of seeds in a fruit (e.g. fig. 4h-h')- 

Although of the living species only 20% have curved seeds the vast majority of the individual 
living plants have curved seeds; so it seems that this probably recent trend in the evolution of the 
genus was successful although the reason for its origin and advantage of its function remains 
obscure. 

Seedlings. Few observations are made. Lubbock (Contr. Knowl. Seedlings, 1892, 206-208, 
fig. 509) noted for S. paniculata (sect. Hopea) : the endocarp does not burst during germination ; 
the radicle emerges by a small hole at the apical narrow end; the hypocotyle elongates, becoming 
curved, finally straightening, carrying up the endocarp containing the embryo. As the cotyledons 
elongate, they push out at the small hole in the endocarp (so to say throw the latter off), and 
finally get free and spread out to the light; they enlarge but remain narrow. The first two leaves 
are opposite, hairy on both sides and serrulate which may persist in leaves of saplings (fig. 
41). 

Spot-characters. In the herbarium a Symplocos of subg. Hopea can mostly easily be spotted by 
spiral, exstipulate, eglandular, serrate or crenate leaves discolouring pale greenish or yellowish or 
greenish-brown, a feature connected with a high Al-content of the tissues. At a very young stage, 
the just expanding leaves have proportionally conspicuous gland-like teeth on the margin. A 
significant character is that in the herbarium the midrib is always sulcate above, with the exception 
of 4 spp. in which it is prominent: S. anomala, S. lancifolia, S. lucida, and S. wikstroemifolia. 

The cup-like 3 bracts (of which 2 bracteoles) below the flower (fig. lib) is also characteristic 
as is the inferior ovary and fruit. 

Innovations and newly expanded leaves are in many species a beautiful violet, afterwards 
changing into violet-brown while the drupes are often blue to black-violet, features found in 
many aluminium-accumulating plants (Eurya, Helicia, etc.). 

Anatomy. For general surveys also covering the older literature, see Solereder, Syst. Anat. 
Dicot. Stuttgart (1899) 587-589 (under Styracaceae) and ibid. (1908) 208-210; Metcalfe & 
Chalk, Anat. Dicot. Oxford (1950) 890-893. Selected references: Janssonius, Mikr. 4 (1925) 
471-498 (wood anatomy); Den Berger, Determinatietabel Malesie, Veenman, Wageningen 
(1949) (wood identification); Janssonius, Blumea 6 (1950) 422^423 & 424 (wood anatomical 
affinities); Desch, Mai. For. Rec. 15 (1954) 591-593 (wood); Zahur, Mem. Cornell Univ. 
Agric. Exp. Stn. 358 (1959) 35 (bark anatomy); Huber, Mitt. Bot. Staatssamml. Miinchen 5 
(1963) 1-48; Baas, Blumea 21 (1973) 201-216 (ecological wood anatomy); Nooteboom, Leid. 
Bot. Ser. 1 (1975) 20-22 (leaf and wood anatomy). 

The wood is characterized by the following primitive set of characters: Vessels solitary and 
with many-barred scalariform perforations. Fibre-tracheids with conspicuously bordered pits 
on both radial and tangential walls. Parenchyma diffuse or dilfuse-in-aggregates. Rays hetero- 
geneous, usually of two distinct si/es. The bark is also of a primitive type with compound sieve 
plates Mechanical hark tissue is poorly developed and composed of groups of sclcreids (Zahur, 
l.< ) The leaf anatomy exhibits few constant characters such as paracytic stoniata, clustered 
crystals and dorsivcntral mcsophyll. Presence Of absence of a hypodcrmis, o\' idiohlaslic leaf 
sclcreids, of a complex vasctilation pattern m the midrib, and of an indumentum varies. The 
diagnostic and systematic value of these characters lemams to be assessed 

T he anatomical e\ulence is inconclusive with respect to a positive indication of the closest 
affinities of Symplocaceae, I lie traditional treatment of the family as a member of the l.lunnlc\ 



212 Flora Malesiana [ser. I, vol. 8 2 

close to Styracaceae must, however, be refuted. The anatomy is more compatible with suggestions 
of a Cornalean or a Thealean alliance as advocated by Nooteboom I.e. 

Palynology. The palynology of the Old World spp. was examined by R. van der Meijden 
(Pollen et Spores 12, 1970, 513-551, 1971, suppl. in my Monograph, 1975, 9-15). The essential 
results are the following: the two main pollen types coincide with the distinction of the two sub- 
genera. In subg. Symplocos there are two minor types, one belonging to the Old World spp., the 
other to those of America. 

In subg. Hopea there are 9 subtypes, but none is apparently peculiar to American spp. The 
distribution of these subtypes is rather complicated and leads to the view of reticulate relation- 
ship, which agrees with the impression gained from macromorphology. Another feature is that 
within the variable species several subtypes are represented, and furthermore that a number of 
subtypes are found in species which are taxonomically not closely related. There is no agreement 
between the shape of the embryo, straight or curved, and pollen subtypes. Echinate pollen is 
found in the Philippine S. whitfordii and in the East Malesian and Pacific varieties of S. cochin- 
chinensis ssp. leptophylla; also the 9 endemic spp. of Symplocos in New Caledonia which are all 
related to this subspecies have echinate pollen. 

In several taxa a certain amount of pollen is sterile and I have ascribed this to hybridization. 

Phytochemistry. Many species of Symplocos, especially from subg. Hopea, contain alu- 
minium compounds, a feature which manifestates itself in the yellow colour of dried leaves. 
Especially when the plants are dried after having been conserved in alcohol vapour according to 
the Schweinfurth method, the yellow colour becomes very intense. The yellow colour is the result 
of a reaction of aluminium compounds with flavonols in the drying leaf. The amounts recorded in 
literature vary between 0.05 and 4.2% of dry weight of the leaves; barks may contain similar 
amounts of aluminium (Chenery, Kew Bull. 1948, 173-183; Analyst, 1948, 501; Nooteboom, 
Leid. Bot. Ser. 1, 1975, 19). Radlkofer (Ber. Deut. Bot. Ges. 22, 1904, 216-224) already men- 
tioned that the ash of Symplocos leaves contains c. 50% aluminium oxide. He also described the 
so-called "Tonerdekorper" in the leaves of Symplocos. These are masses of colourless material 
filling often large parts of the cells, predominantly in the palissade parenchyma. According to 
Radlkofer these masses consist mainly of aluminium compounds. Kratzmann (Sitz. Ber. Ak. 
Wiss. Wien, 1913, 31 1-336) found that these aluminium bodies also contain much other material, 
for instance silicates, and that the aluminium is also accumulated in other parts of the leaf. Neger 
(Flora N.F. 16, 1923, 326-330) observed that the development of plants of Symplocos lucida 
(Thunb.) S. & Z. depends on the amount of aluminium compound in the solution they are culti- 
vated on. Plants grew best on a solution containing 1 promille aluminium. Besides aluminium 
many other compounds are found (Hegnauer, Chemotaxonomy der Pflanzen 6, 1973). The 
more important are: 1) Phenolic compounds (see also Bate Smith, J. Linn. Soc. Bot. 58, 1952, 
95-173). Gallic and ellagic acid seem to be rather common. Leucoanthocyanins occur in varying 
amount. Bate Smith I.e. also found quercetin, and caffeic acid. In the bark of S. lucida (Thunb.) 
S. & Z. the lignan glycoside symplocosin has been found, and traces of methylsalicilate were 
demonstrated in the bark of several species. True tannins were not yet found in Symplocos. 
2) Alkaloids. Only for two species structurally known alkaloids were described. More research is 
needed. 3) Saponins. In several species saponin-like compounds were found, as well in the bark 
as in the leaves. — R. Hegnauer. 

Chromosomes. In my monograph I have given an account of chromosome numbers, which 
are unfortunately too few. However, the majority is n = 11, with some deviations; rarely 2n = 
24, and one count of the North American S. tinctoria of In = 14, all in subg. Hopea. The one 
count known of subg. Symplocos in Malesia yielded 2n = c. 90 (2n = 88 would fit an octoploid). 
It would be too rash to conclude that polyploidy would be normal in that subgenus. 

On the other hand it may tentatively be concluded that species in subg. Hopea are diploid, with 
x = 11. This does not fit the numbers found in other Ebenales families, nor in Theaceae, but it 
does agree with Cornaceae sens. lat. 

Uses. As timber Symplocos has no great value, according to Heyne (Nutt. PI. 1927, 1262). 
Leaves and bark of Symplocos contain a fair amount of alum, both in Asian and American spp. 
(cf. Ber. Deut. Bot. Ges. 22, 1904, 126). This was commonly used, mostly from decoctions of the 
bark, in dyeing processes (red and brown), e.g. in the batik industry in Java. Rumphius already 



1977] 



Symplocaceae (Nooteboom) 



213 



mentioned this use from the Moluccas. Several species were used for this purpose, e.g. S. 
cochinchinensis, S. faseiculata, S. odoratissima (Heyne, I.e.). The same compound is probably 
also the constituent active in medicinal uses against so-called sprue ('thrush') as 'obat seriawan. 

Notes. Identification of material of Symplocos is for several reasons far from easy. Because of 
simultaneous flowering flowers and fruits are practically never found together and both stages 
are properly needed. Only few species possess well definable vegetative characters. Moreover, a 
few widespread species have proved to be rather variable, to a fair degree by racial differentiation. 
These are the reasons that besides a general key in which all characters are used, I have found it 
useful to add a number of partial keys for islands or island groups in a double series, either for 
fruiting or for flowering material. 

In fig. 5 a scheme is given elucidating the way in which for this genus descriptive terminology is 
used in the keys and descriptions. 






Fig. 5. Schemes elucidating descriptive terminology used in the text. — A: a length of leaf, b width of 

leaf, c divided by b is leaf index, d length of acumen. — B: way of expressing base angle a. — C: deflorated 

flower; br bract, bra bracteole, c length of calyx, cl length of calyx lobes, ov height of ovary. 



KEY TO THE SUBGENERA 

1. Petals connate at least halfway up. Leaves usually not becoming yellow when drying, not discolouring, 
spirally arranged, entire. Flowers very fragrant. Seeds and embryo straight. Spp. 1-2 

1. subg. Symplocos 

I. Petals connate only at the very base. Leaves usually becoming more or less yellow or greenish yellow 
when dried. Leaves spirally arranged or distichous, exceptionally in pseudowhorls. Flowers not or 
mostly only faintly fragrant. Seeds and embryo straight or curved. Spp. 3 58 .... 2. subg. Hopca 



1. Subgenus S> mplocos 

Cf. Noot. Lcid. Bot. Ser. 1 (1975) 36. -- Cordylobkute MOR. Bot /at. 6 (1848) 

606; Rn>] . I I. Mai. Pen. 2(1923)307; Alston, Handb. 1 1. Ceyl. 6(Suppl.)(1931) 

Symplocos feet. Cordylobkute B. & H. Gen. PL 2 (1876) 669; Brand, Pfl. 

R. Heft 6 (1901) 88; Sins. Hull. Bot. Gaid. BtZg III. 17(1948)429. Symplocos 

wbg. Cordylobkute Gamble, J. As. See. Beng. 74, ii (1906) 24s. I i^. 6. 

Leaves usually not becoming yellov. v. hen dry. Corolla tubular, ereet, often tO 
above the middle adherent to the staminal tube and then Hidden!) expanded; 



214 



Flora Malesiana 



[ser. I, vol. 8 2 



margins of the petals free, thus sometimes obscuring the coalescence. Stamens 
monadelphous; free part of filaments ribbon-shaped, in several whorls, in the outer 
whorl often very short, always suddenly attenuate below the anther. Fruits 2-5- 
celled, usually none of the cells aborted. Seeds straight, cylindrical. 

Distr. Tropics of Indo-Malesia and South America, largely within 30° N and S, more than 100 spp. 
described from the New World, in Malesia 2 spp. Fig. 1 . 
Ecol. Rain-forest, from the lowland up to c. 3300 m (Mt Kinabalu). 



KEY TO THE SPECIES 



1. Calyx c. 6(-10) mm long. Corolla 2 1 l 2 -5 cm long. Fruits 3-5 cm long 
1. Calyx 3-5 mm long. Corolla V2-l 3 /4 cm long. Fruits l-l'/a cm long 



1. S. henschelii 
2. S. pendula 



1. Symplocos henschelii (Mor.) Bth. ex Clarke, 
Fl. Br. Ind. 3 (1882) 588, quoad nomen et basionym, 
excl. stirp.; Brand, Pfl. R. Heft 6 (1901) 89; Bull. 
Herb. Boiss. II, 6 (1906) 750; Koord. Atlas 2 (1914) 
t. 390; Steen. Bull. Bot. Gard. Btzg III, 17 (1948) 
440, f. 2 a-1 ; Nova Guinea n.s. 10 (1959) 210; Back. 
& Bakh./. Fl. Java 2 (1965) 204; Steen. Mt. Fl. 
Java (1972) pi. 52-3; Noot. Leid. Bot. Ser. 1 
(1975) 37, pi. lg. — Cordyloblaste henscheli Mor. 
Bot. Zeit. 6 (1848) 606. — Eugeniodes henscheli 
O. K. Rev. Gen. PI. 2 (1891) 975. — 5. nageli 
K. & V. Bijdr. 7 (1900) 159. — S. scortechinii King 
& Gamble, J. As. Soc. Beng. 74, ii (1906) 250. — 
Cordyloblaste scortechinii Ridl. Fl. Mai. Pen. 2 
(1923) 309. — S. dolichantha Merr. Sar. Mus. J. 3 
(1928) 545. — S. stenosepala Steen. Bull. Bot. 
Gard. Btzg III, 17 (1948) 444, f. 2 m-n. — Fig.6a-c. 

For further synonyms see under the variety. 

Shrub, or mostly a tree, to 30 m; innovations 
glabrous to grey or rusty velvety. Leaves glabrous, 
sometimes the midrib above and underside hairy, 
7-17(-22) by 3-7V 2 cm; petiole l l r l 1 UL-2) cm. 
Racemes up to 10 cm, incl. bracts and flowers grey 
or rusty tomentose, short-peduncled, 1—12- 
flowered. Bracts narrow-triangular; pedicels 0-6 
mm, with 2(-3) tiny bracteoles. Calyx lobes 
rounded to triangular, mostly erect, 1-4V 2 by 
2-3 mm, persistent. Corolla sericeous (in Mai.), 
club-shaped in bud, 2V2-5 cm, connate for 3 / 5 - 3 / 4 , 
tube 3-4 mm 0, lobes spathulate. Staminal tube 
1 I 2 cm shorter than corolla, adnate to the corolla 
tube except towards apex, free part 1 / 2 -l l l2cm; 
anthers 20-110, filaments unequal. Ovary 3-4- 
celled; ovules 2-4 per cell, usually only 1 develop- 
ing. Fruit obovoid to spindle-shaped, 3-5 by 2-3 
cm; mesocarp thick, hard-fleshy to ± woody. 

Distr. Continental SE. Asia (Burma, Thailand, 
Indo-China) and West Malesia (Sumatra, Malay 
Peninsula, W. Java, Borneo), a distinct subspecies 
in Thailand. 

Note. Additional material has shown that 
S. stenosepala Steen. cannot be upheld and, more- 
over, that 5. maingayi Clarke deserves only 
varietal rank. 



KEY TO THE VARIETIES 

1. Leaves and twig ends usually glabrous. Free 

part of staminal tube 7-15 mm. 

a. var. henschelii 
1. Twig ends and leaves underneath hairy. Free 

part of staminal tube 5-7 mm b. var. maingayi 



a. var. henschelii. — Fig. 6a-b. 

Shrub or tree, up to 25 m, 45 cm 0. Twigs glab- 
rous, the youngest ones sometimes more or less 
grey or rufescent appressedly pubescent to velvety 
or tomentose. Leaves glabrous, or the midrib 
beneath sparsely short fine-hairy, rarely with same 
indument as var. maingayi. Free part of staminal 
tube 7-15 mm; anthers (40-)55-75(-110), in the 
upper 5-10 mm, ascendent and nearly sessile 
above to descendant on a slender filament below, 
the lowest ones hanging from a 2-5 mm long fila- 
ment. Fruit with ± fleshy mesocarp. 

Distr. As the species. 

Ecol. Below 1 100 m in mixed dipterocarp forest, 
also once in swamp forest, and on podsol (Kalabit), 
at higher altitude in oak-chestnut mountain forest, 
also on ridges and in mossy forest, 600-2000 m (in 
continental SE. Asia at 130-800 m). Fl. Jan.-Dec., 
fr. Febr.-Sept. 

Vern. Sumatra: kayu djaram-djaram bosi, 
Batak; Borneo: te baradang, Sarawak, Kalabit, 
yum, Kenyah lang., lamau-lamau, Brunei. 

b. var. maingayi (Clarke) Noot. Leid. Bot. Ser. 1 
(1975) 39. — S. maingayi Bth. ex Clarke, Fl. Br. 
Ind. 3 (1882) 588; Brand, Pfl. R. Heft 6 (1901) 90; 
K. & G. J. As. Soc. Beng. 72, ii (1906) 249; Steen. 
Bull. Bot. Gard. Btzg III, 17 (1948) 445. — Euge- 
niodes maingayi O. K. Rev. Gen. PI. 2 (1891) 975. 
— Cordyloblaste maingayi Ridl. Fl. Mai. Pen. 2 
(1923)309. — Fig. 6c. 

Tree up to 21 m, 40 cm 0. Twigs densely rusty 
tomentose or velvety, glabrescent. Leaves sparsely 
fine-hairy beneath, especially on midrib and nerves, 
to greyish tomentose or velvety. Free part of 
staminal tube 5-7 mm; anthers 20-60, in the upper 
5 mm, on a very short (V4-V2 mrn ) tr, in free part of 
the filaments. Fruit with ± woody mesocarp. 

Distr. Malesia: Malay Peninsula and Borneo 
(Sarawak, Brunei). 

Ecol. Evergreen primary and depleted lowland 
forest, 15-150 m; in Borneo often on low sandy 
ridges, raised beaches, and large sandy podsols 
(kerangas). Fl. April-May, fr. Jan. 

2. Symplocos pendula Wight, Ic. 4 (1848) 10, 
t. 1237; 111. Ind. Bot. 2 (1850) t. 151-b, 7-12; 
Clarke, Fl. Br. Ind. 3 (1882) 587; Brand, Pfl. R. 
Heft 6 (1901) 88; Steen. Bull. Bot. Gard. Btzg III, 
17 (1948) 437; Noot. Leid. Bot. Ser. 1 (1975) 40, 
pi. lh. — S. scortechinii (non K. & G.) Ridl. J. 
Linn. Soc. Bot. 38 (1908) 315. — S. pulcherrima 



1977] 



Symplocaceae (Nooteboom) 



215 







Fig. 6. Symplocos henschelii (Mor.) Bth. ex Clarke var. henschelii. a. Flower, b. fruit, exocarp halved. — 
S. henschelii var. maingayi (Clarke) Noot. c. Fruit. — 5. pendula Wight var. pendula. d. Flower. — 
S. pendula var. hirtistylis (Clarke) Noot. e. Fruit. All nat. size (a Wilson 2547, b after Steen. 1972, pi. 
52-3b, c Kostermans 9328, d father Anglade s.n., e Meijer 3618). 



RrDL. J. Fed. Mai. St. Mus. 6 (1915) 160. — 
Cordyloblaste pulcherrima Ridl. Fl. Mai. Pen. 2 
(1923)308. — Fig. 6d-e. 

For further synonyms see under the variety. 

var. pendula. — Fig. 6d. 

Small shrub ' 2 -3 m or tree up to 27 m and 50 cm 
0. Twigs glabrous to rusty tomentose. Leaves 
glabrous or nearly so, elliptic to obovate or orbicu- 
lar, entire to crenate, apex rounded to acuminate, 
(1-)2 1 2 -12' 2 by (l-)l" 2 -6cm; nerves 4-8(-ll) 
pairs; petiole (l-)5-15mm. Racemes very short, 
sometimes flowers solitary. Bracts to 1 mm. Brac- 
teoles 2-4, narrow-triangular. Pedicels 0-5 mm, 
longer in solitary flowers. Calyx lobes very short 
and rounded, ciliate. Corolla tubular-trumpet- 
shaped, (5-)10-17mm, fleshy, silver-white to 
creamy, fragrant, the petals connate halfway up, 
spathulate, rounded at apex, glabrous to tomentose. 
Staminal tube adnate to corolla except for upper 
3-5 mm, hairy to glabrous inside; anthers 30-50 
(-80). Ovary semi-inferior, glabrous, the apex 
semi-globose, c. l*/j mm high, densely grey-hairy; 
style c. 1 cm, more or less hairy at the base to 
glabrous at the apex. Fruits spindle-shaped, 10-15 
by 3-6 cm, green pinkish red, the enlarged calyx 
lobes surrounding the hairy, conical, persistent 
style-base. 

Distr. Continental SE. Asia (Ceylon, Deccan, 
Hainan), in Malesia: Malay Peninsula. 

Ecol. Mountain forests and open heath and 
scrub, often on ridges, 600-1750 m. Fl. Febr.- 
March,/r. Sept. 

var. hirtistylis (Q arku Noot. Leid. Bot. Scr. I 



(1975) 42, f. 2a, with full synonymy. — S. henschelii 
(non Bth.) Clarke, Fl. Br. Ind. 3 (1882) 588, pro 
stirp., incl. var. hirtistylis Clarke. — S. confusa 
Brand, Pfl. R. Heft 6 (1901) 88; Bull. Herb. Boiss. 
II, 6 (1906) 750; K. & G. J. As. Soc. Beng. 74, ii 
(1906) 248; Brand, Philip. J. Sc. 3 (190") Bot. 3; 
Merr. En. Philip. 3 (1923) 297; Steek J. Am. 
Arb. 28 (1947) 423; Bull. Bot. Gard. Btzg III, 17 
(1948) 432. — S. albifrons Brand, Pfl. R. Heft 6 
(1901 ) 88 ; Bull. Herb. Boiss. II, 6 (1906) 750 ; Nova 
Guinea 14 (1924) 189. — 5. capitellata Brand, 
Pfl. R. Heft 6 (1901) 88; Bull. Herb. Boiss. II, 6 
(1906) 750; Nova Guinea 14 (1924) 188. — S.fox- 
worthyi Brand, Philip. J. Sc. 3 (1908) Bot. 3; 
Merr. En. Philip. 3 (1923) 299. — Styrax obovatus 
Ridl. J. Str. Br. R. As. Soc. n. 61 (1912) 8. — 
S. obovata Ridl. J. Fed. Mai. St. Mus. 6 (1915) 51. 
— S. crenulata Ridl. I.e. — S. novoguineensis 
Gibbs, Arfak (1917) 176. — Cordyloblaste obovata 
Ridl. Fl. Mai. Pen. 2 (1923) 308. — Cordyloblaste 
crenulata Ridl. I.e. 309. — S. atrata Brand, 
Nova Guinea 14 (1924) 188. — 5. topica Brand, 
I.e. 189. — Fig. 6e. 

Ovary hairy. 

Distr. Continental SE. Asia (N. Burma, Indo- 
China, China, Japan, Formosa), throughout 
Malesia, except Java and Lesser Sunda Is. 

Ecol. Primary and secondary montane and sub- 
alpine forest, mossy forest, often common on 
ridges, or in open fern thickets (Tamrau), on sand 
or clay, 1500 3300 m, but in kerangas forest in 
Sarawak at 800 m. Fl. March-Aug. (Sept.-Febr.), 
I> . \ ebr. April, July Sept. 

At higher altitude often a dwarf shrub with small 
leaves, but sometimes also a dwarf shrub with large 
leaves in high forest. 



2. Subgenus Ho pea 

Clarke, Fl. Br. Ind. 3 (1882) 572; Brand, Pfl. R. Heft 6 (1901) 25; Noot. Leid. 
Bot. Scr. 1 (1975)43, with full synonymy. — Hopea Linne, Mant. (1767) 105, nom. 



216 



Flora Malesiana 



[ser. I, vol. 8 2 




O#50 



II lp 



8 9 10 10 11 a 12 



o 'OiooOi 



16-1C 16 id 16 2e 16 3f 16 4g 16 4g 




16-4g 16 4g 16 4g 16 4h 16-4J 16-41 16 4m 16 - 4m 16 4fl 16-40 







i f I i 

16-4P 16-4q 16-4T 164T 16 4S 16 4t 16 4U 16 4U 16 4W 



16 4W 16 4X 17 



20 C 



20 d 20 f 21 22 23 






w iff yuo 



24 27 28 30 31 32 33a 33e 34 35 









46 47a 47 b 48 50 51 52 53 55 56 57 58 

Fig. 7. Fruits in outline, in the dried state. Of each fruit the voucher specimen is cited by the number of the 
taxon. If for showing variability more fruits of the same taxon are drawn, read from left to right corres- 
ponding with the voucher numbers. All drawings natural size. — 3 Chew Wee Lek 938 — 3 CF 104879 — 
4 SAN 56690 — 5 de Wilde 13773 — 6 A. Ernst 736 — 7 King's Coll. 6179 — 8 Meijer 7581 — 9 



1977] Symplocaceae (Nooteboom) 217 

rejic. — Dicalix Lour. Fl. Coch. 1 (1790) 663; Bl. Bijdr. (1826) 1116 {'Dicalyx 1 '). — 
Sahara Reinw. Syll. Ratisb. 2 (1825) 12. — Carlea Pr. Epim. Bot. (1851) 216. — 
Baranda LLanos, Mem. Ac. Cienz. Madrid 3, 2 (1857) 502. — Eugeniodes O. K. 
Rev. Gen. PI. 2 (1891) 409, 975, nom. illeg. — Fig. 7-20. 

Leaves usually becoming more or less yellow when drying. Petals glabrous, or 
hairy in only few species, connate only at the very base, mostly expanded. Stamens 
monadelphous to pentadelphous, only connate at the very base (for at most 2 mm) ; 
filaments cylindrical, slender to rather stiff, often gradually attenuate towards the 
anther. Fruits 2-3(-5?)-celled, often 1 -celled by abortion. Seeds either straight or 
curved, and then with curved embryo. 

Distr. About 150 spp., as for the genus. Fig. 1. 

Note. As explained in the note under the genus, a general overall key is given to all species, as much as 
possible based on vegetative characters and on flowering material. 

To facilitate identification additional local keys are given for the main Malesian islands or island groups, 
one each for flowering and for fruiting material. 

In addition in fig. 7 fruits are drawn of all species as far as available in the dried state. They have been 
numbered according to the number of the taxa. The following terminology has been adopted for fruit- 
shapes : 

globose 34, 41b ampulliform 23, 42c, 43 

ellipsoid 4, 19, 33e, 46 spindle-shaped 38: fig. 19d. 

ovoid 48 cylindrical 20c, 21 

obovoid la, lb: fig. 6c 

It should be observed that the shape of the stone may differ from the shape of the fruit and that for 
instance ovoid fruits may possess an ampulliform stone. 

There is no strict relation between the shape of the seed and the shape of the fruit or stone, but ampulli- 
form fruits have always a curved seed and curved embryo and spindle-shaped and cylindrical fruits have 
always a straight seed and embryo. 

Besides the overall-shape of the drupe, the shape of the stone can be important : sometimes it bears 
lower or higher ridges, which ornamentation provides good characters. 



KEY TO THE SPECIES 

1. Leaves (pseudo-)vertici Hate. 
2. Upper side of leaves glabrous. Twigs hirsute 55. S. verticillifolia 

2. Upper side of leaves hairy. Twigs tomentose 30. S. herzogii 

1. Leaves not verticillate. 

3. Midrib prominent on the upper surface. 
4. Twigs glabrous. 

5. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 

37. S. wikstroemifolia 

5. Leaves evenly distributed, glabrous 35. S. lucida 

4. Twigs hairy. 

Hai mik/. 2197 — 10 Clemens 32525 — 10 Clemens 32478 — 11a bb 23324— 12 SAN 46543 — 14 
K0BTERMANS9158 -• 15a Kosmkmans & Anta 527 — 16-la Forbes 861 — 16-lblvtEUEK 1690- 16 lc 
CLEMENS l 7 224— 16-ld BS4476 16 2c LarSEN C.S. 887— 16-3f NGF 33643— 16-4g ANU 2027 
16-4g A.( . Smiih 1054— 16-4gBW4';7<) l6-4g Giimsimi 3918 16 4g NGh 28481 — 16-4h Vink 
17308 16 4i Brass 28343 16 41 BRASS 29919 - 16-4m LeDBRMANN 8 l >46 16 4m T. G. HARTLEY 
13135 — 16 4n PULLEN 479 16 4o NGF 49168 16 4p l'i iiin 7783 16 4q KOSTERMANS & 

Wirawan 878 l6-4r Nicolas 19 l6-4r van Baloooy 862 16-4* Kalkman 5128 16 4t Vink 

16079 -- 16 4u KOBTERMANS 2375 16 4u FORBES P. P. 652 — 16 4vv Brass 28191 16 4\s CLEMENS 

1661- I', k NGF 23728 17 Clemens 33706 19 Koorders 15596 20c van Beusbkom c*. 837 
20d f NDI i< i 2580 201 < I 97832 21 bb 22503 22 Ja( OM 5766 23 SAN A2240 - 24 Mi rrii I 

6148 — 27 bs 45592 - 28 KEP/FRI 8236 K> T. G. Hartley 12509 11 Nooteboom & Aran 1500 

32 HlLDEBRAND 55 33fl SAN 65017 3 V: SAN 44386 - 34 I'NII 18 483 35 BURGER J.ft. 36 

( cl. II 374 37 I C. HOW 73506 40 liS 26447 41a Ja 7723 41b SAN 57045 — 42 la BURN 

Murdoch 340 42-3f Dino Hou 274 42-3f Nooteboom 2229 42 3gHSX3753 42 2c ( T 98890 

— 43 Kaiewski 1208 44 S 17287 45 Carr 12782 46 Ridley 16102 47b Robinson & Kloss 199 

-47bMEUER7665 48Ismabl9 50 Beccari P. S. 106 51Koelz29538 52 Clemens 32559 53 

S26305 — 55 I'M I 14397 56 US 45675 or 45775 - 57 Jacobs 7484 58 Noon BOOM 1491. 



218 Flora Malesiana [ser. I, vol. 8 2 

6. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 

37. S. wikstroemifolia 
6. Leaves evenly distributed, glabrous or sparsely fine hairy beneath. 

7. Underside of leaves glabrous. Corolla 4-6 mm 4. S. anomala 

7. Underside of leaves hairy. Corolla 2' 2 -4 mm 34. S. lancifolia 

3. Midrib sulcate above. 

8. Corolla (densely) hairy 41. S. odoratissima 

8. Corolla glabrous. 
9. Twigs hairy. 
10. Underside of leaves glabrous. (When petiole and leaf margin beset with closely spaced vesicular 
glands: 3. S. adenophylla). 

11. Leaves distichous 33. S. laeteviridis 

11. Leaves spirally arranged. 
12. Calyx and ovary glabrous. 

13. Petiole 0-5 mm 16-4. S. cochinchinensis ssp. leptophylla 

13. Petiole more than 5 mm. 

14. Leaves shorter than 5 cm 16-4. S. cochinchinensis ssp. leptophylla 

14. Leaves longer than 5 cm. 

15. Disk hairy 16-4. S. cochinchinensis ssp. leptophylla 

15. Disk glabrous. 
16. Twigs (appressedly) pubescent, puberulous or pilose. Seeds not straight. 

16. S. cochinchinensis 
16. Twigs tomentose or tomentellous. 
17. Petiole 12-17 mm. Acumen 2-7 mm long. Nerves 8-12 pairs. Fruits more than 10 mm 

long 5. S. atjehensis 

17. Petiole 5-12 mm. Acumen longer than 7 mm. Nerves 10-16 pairs. Fruits to c. 10mm long. 

28. S. glomerata 
12. Calyx and or ovary hairy. 
18. Leaves crowded towards the end of the twigs, the latter tapering off towards the apex. 

44. S. polyandra 
18. Leaves evenly distributed, twigs not obviously tapering off. 
19. Ovary glabrous. 

20. Inflorescence only 1 -flowered 38. S. multibracteata 

20. Inflorescence more-flowered. 

21. Disk hairy 16-4. S. cochinchinensis ssp. leptophylla 

21. Disk glabrous. 
22. Seed and embryo uncinately curved towards the base 

16-4. S. cochinchinensis ssp. leptophylla 
22. Seed and embryo twice curved .... 16-1. S. cochinchinensis ssp. cochinchinensis 
19. Ovary hairy. 

23. Calyx glabrous 16-4. S. cochinchinensis ssp. leptophylla 

23. Calyx hairy. 
24. Bracts caducous. 
25. Inflorescence an often branched raceme to 4 cm. Calyx 1-2 mm long 47. S. robinsonii 

25. Inflorescence a 1-3-flowered short spike. Calyx c. 3 mm .... 10. S. brachybotrys 
24. Bracts persistent. 

26. Petiole 0-5 mm 16-4. S. cochinchinensis ssp. leptophylla 

26. Petiole more than 5 mm. 

27. Seeds straight 24. S. filipes 

27. Seeds not straight 16-4. S. cochinchinensis ssp. leptophylla 

10. Underside of leaves hairy. 
28. Leaves distichous. 
29. Nerves up to 6 pairs. 

30. Angle of leaf base more than 90° 31. S. johniana 

30. Angle of leaf base less than 90°. 

31. Disk glabrous 52. S. trichomarginalis 

3 1 . Disk hairy 33. S. laeteviridis 

29. Nerves (5-)6 pairs or more. 
32. Leaves longer than 5 cm (mean length). 
33. Flowers c. 3 in an up to 3 cm long lax raceme. Fruits 10-14 mm long. Stamens c. 90 or more. 

17. S. colombonensis 
33. Inflorescence usually different. Fruits to c. 12 mm long. Stamens c. 70 or less. 
34. Inflorescence a fascicle. Bracts to c. 1 mm long, persistent, bracteoles persistent. Ovary 
c . 1 mm high, calyx c. 1 mm long, lobes not becoming longer by tearing. Corolla c. l-A 1 ^ 

mm. Style base hairy. Fruits ampulliform 23. S. fasciculata 

34. Inflorescence not a fascicle. Bracts longer than 1 mm, caducous, bracteoles caducous. 
Ovary more than 1 mm high, calyx longer than 1 mm, lobes becoming longer by tearing. 
Corolla more than 4 mm long. Style base glabrous. Fruits ovoid to ellipsoid 

33. S. laeteviridis 



1977] Symplocaceae (Nooteboom) 219 

32. Leaves shorter than 5 cm. 
35. Inflorescence only 1 -flowered. 
36. Reticulation not prominent. Ovary c. 1 mm high, calyx longer than 2 mm, lobes c. 3 mm. 
Bracts several. Corolla c. 4 mm. Fruits 8-9 mm long .... 52. S. trichomarginalis 

36. Reticulation present beneath. Ovary 1-1 ' 2 mm high, calyx c. 2 mm long, lobes 1-1 ' 2 mm 
long. Bract 1. Corolla 4-6 mm long. Fruits 10-12 mm long .... 58. S. zizyphoides 

35. Inflorescence more-flowered. 

37. Bracts and bracteoles caducous 33. S. laeteviridis 

37. Bracts and bracteoles persistent. 

38. Disk hairy. Fruits c. 10 by 5 mm 22. S. deflexa 

38. Disk glabrous. Fruits 10-12 by 5-6 mm 58. S. zizyphoides 

28. Leaves spirally arranged. 
39. Upper side of leaves hairy. 

40. Angle of leaf base more than 90 D 13. S. calycodactylos 

40. Angle of leaf base less than 90". 

41. Leaf margin (and petiole) beset with closely spaced glands 3. S. adenophylla 

41. Leaf margin (and petiole) often glandular but glands not closely spaced. 
42. Ovary hairy. Fr. cylindrical, 13-18 by 3-5 mm. Embryo straight . . . . 20. S. crassipes 
42. Ovary glabrous. Fr. ampulliform, 6 by 4 mm. Embryo twice curved 43. S. paucistaminea 
39. Upper side of leaves glabrous. 
43. Calyx and ovary glabrous. 
44. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 

37. S. wikstroemifolia 
44. Leaves evenly distributed, glabrous or longer hairs beneath. 
45. Seeds straight. 

46. Leaf index 2-3. Fruits less than 20 mm long 5. S. atjehensis 

46. Leaf index more than 3. Fruits longer than 20 mm. 

47. Nerves less than 10 pairs 15. S. cerasifolia 

47. Nerves more than 10 pairs 15b. S. cerasifolia var. grandifolia 

45. Seeds not straight 16-4. S. cochinchinensis ssp. leptophylla 

43. Calyx and or ovary hairy. 

48. Leaves crowded towards the end of the twigs 37. S. wikstroemifolia 

48. Leaves evenly distributed. 
49. Ovary glabrous. 
50. Disk hairy. 

51. Inflorescence only 1 -flowered. Seeds straight 38. S. multibracteata 

51. Inflorescence more-flowered. Seeds not straight. 

52. Petiole 0-5 mm 34. S. lancifolia 

52. Petiole more than 5 mm 16-4. S. cochinchinensis ssp. leptophylla 

50. Disk glabrous. 

53. Ovary to c. 1 mm high 16-1. S. cochinchinensis ssp. cochinchinensis 

53. Ovary more than 1 mm high. 
54. Twigs (appressedly) pubescent, puberulous or pilose. 

16-4. S. cochinchinensis ssp. leptophylla 
54. Twigs not appressedly pubescent or puberulous. 

55. Leaf index 2-3. Bracts persistent, shorter than 3 mm, bracteoles persistent. Calyx 
lobes not becoming longer by tearing. Corolla shorter than c. 4 mm. Stamens less than 
30. Fruits to c. 10 mm long. Mesocarp fleshy (shrivelled when dry). Seeds not straight. 

16-4. S. cochinchinensis up. leptophylla 
55. Leaf index more than 3. Bracts caducous, longer than 3 mm, bracteoles caducous. 
Calyx lobes becoming longer by tearing. Corolla c. 5 mm long. Stamens more than 
30. Fruits more than 20 mm long. Mesocarp woody or corky. Seeds straight 

15. S. cerasifolia 
49. Ovary hairy. 

56. Calyx glabrous 16-4. S. cochinchinensis ssp. leptophylla 

56. Calyx hairy. 
57. Bracts caducous. 
58. Leaves longer than 15 cm. 
59. Fruits more than 10 mm long, 2-5-cellcd. Mesocarp woody or corky. Stone with high 
lengthwise not interrupted ridges. Seeds straight . 15b. S. cerasifolia var. grandifolia 

59. Fruits to c. 10 mm long, 1-celled. Mesocarp thin, friable in dry state. Stone with I 
transverse constriction at one side. Seeds not straight 48. S. rubi^inosa 

58. Leaves shorter than c. 15 cm. 

60. Calyx lobes longer than I' , mm. Style base hairy. 

61. Leaves shorter than 5 cm 10. S. brachybotrys 

61. Leaves longer than 5 cm. 

62 Inflorescence a raceme 50. S. sumatrana 

62. Inflorescence a I -3-flowcred short spike or a spike to 4 cm. 



220 Flora Malesiana [ser. I, vol. 8 2 

63. Petiole 3-4 mm. Ovary c. 1 mm high, calyx c. 3 mm, lobes longer than 2 l l 2 mm. 
Ovary (appressedly) pubescent. Disk inconspicuous. Fruits c. 5 mm broad, 

1 -celled. Seeds not straight 10. S. brachybotrys 

63. Petiole 5-7 mm. Ovary c. l l l 2 mm high, calyx 2 mm long, lobes lVr-2Va mm long. 
Ovary sericeous. Disk clearly present. Fruits more than 5 mm broad, 3-celled. 

Seeds straight 50. S. sumatrana 

60. Calyx lobes 1 / 2 — IV2 mm long. Style base glabrous. 

64. Nerves more than 10 pairs 47. S. robinsonii 

64. Nerves less that 10 pairs. 
65. Ovary c. 1 mm high, lobes triangular. . 42-lb. S. ophirensis var. densireticulata 

65. Ovary 1-2 mm high, lobes not triangular 47. S. robinsonii 

57. Bracts persistent. 
66. Inflorescence only 1 -flowered. 

67. Angle of leaf base more than 90° 31. S. johniana 

67. Angle of leaf base less than 90°. 

68. Leaf index 4-7. Ovary c. l'/ 4 mm high. Stamens less than 30. Disk hairy, incon- 
spicuous. Stone ?smooth 49. S. salicioides 

68. Leaf index 1.3-4. Ovary to c. 1 mm high. Stamens more than 30. Disk glabrous, 

clearly present. Stone with ridges or grooves 20. S. crassipes 

66. Inflorescence more-flowered. 

69. Seeds straight. 

70. Bracts to c. 1 mm long. 
71. Ovary to c. 1 mm high. 
72. Reticulation fine. Calyx longer than 1 mm. Inflorescence a much reduced often 
clustered spike. Fruits not ampulliform, 13-18 mm long ... 20. S. crassipes 

72. Reticulation coarse. Calyx c. 1 mm long. Inflorescence a fascicle. Fruits ampulli- 
form, 5-7 mm long 23. S. fasciculata 

71. Ovary more than 1 mm high. 

73. Leaf index more than 3. Calyx lobes longer than 1 / 2 mm . . . 3. S. adenophylla 
73. Leaf index 2-3. Calyx lobes to c. 1 / 2 mm long 24. S. filipes 

70. Bracts longer than 1 mm. 

74. Angle of leaf base more than 90° 20. S. crassipes 

74. Angle of leaf base less than 90°. 

75. Underside of leaves especially hairy on the margin 20. S. crassipes 

75. Underside of leaves not especially hairy on the margin. 
76. Calyx lobes longer than l'^rnm. Disk hairy. Stamens more than 30. Style base 

hairy. Fruits more than 10 mm long 20. S. crassipes 

76. Calyx lobes 1-1 l j 2 mm long. Disk glabrous. Stamens less than 30. Style base 

glabrous. Fruits c. 10 mm long 56. S. vidalii 

69. Seeds not straight. 

77. Disk glabrous 16-4. S. cochinchinensis ssp. leptophylla 

77. Disk hairy. 

78. Seed and embryo U-shaped 34. S. lancifolia 

78. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 
9. Twigs glabrous. 
79. Underside of leaves hairy. 
80. Leaves crowded towards the end of the twigs, minutely appressedly hairy beneath 

37. S. wikstroemifolia 

80. Leaves evenly distributed. 
81. Calyx and ovary glabrous. 

82. Disk hairy 16-4. S. cochinchinensis ssp. leptophylla 

82. Disk glabrous. 

83. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 

83. Seed and embryo not uncinately curved towards the base 5. S. atjehensis 

81. Calyx and/or ovary hairy. 

84. Leaves distichous 33. S. Iaeteviridis 

84. Leaves spirally arranged. 

85. Leaves shorter than 5 cm 10. S. brachybotrys 

85. Leaves longer than 5 cm. 
86. Calyx glabrous. 
87. Calyx lobes becoming longer by tearing. Seeds straight 20. S. crassipes 

87. Calyx lobes not becoming longer by tearing. Seeds not straight. 

16-4. S. cochinchinensis ssp. leptophylla 
86. Calyx hairy. 

88. Petiole 3-4 mm 10. S. brachybotrys 

88. Petiole more than 5 mm. 

89. Ovary glabrous 16-4. S. cochinchinensis ssp. leptophylla 



1977] Symplocaceae (Nooteboom) 221 

89. Ovary hairy. 

90. Bracts and bracteoles caducous 48. S. rubiginosa 

90. Bracts and bracteoles persistent. 

91. Bracts to c. 1 mm long. Seeds straight 24. S. filipes 

91. Bracts longer than 1 mm. Seeds not straight. 

16-4. S. cochinchinensis ssp. leptophylla 
79. Underside of leaves glabrous. 

92. Leaves distichous 33. S. laeteviridis 

92. Leaves spirally arranged. 
93. Calyx and or ovary hairy. 
94. Ovary glabrous. 
95. Bracts caducous. 

96. Leaves shorter than 5 cm 12. S. buxifolia 

96. Leaves 9-15 cm 21. S. cylindracea 

96. Leaves longer than 15 cm 7. S. barringtoniifolia 

95. Bracts persistent. 

97. Inflorescence only 1 -flowered 12. S. buxifolia 

97. Inflorescence more-flowered. 

98. Disk hairy 16-4. S. cochinchinensis ssp. leptophylla 

98. Disk glabrous. 

99. Calyx lobes to c. l /a mm long 16-4. S. cochinchinensis ssp. leptophylla 

99. Calyx lobes longer than l /a mm - 

100. Ovary to c. 1 mm high 16. S. cochinchinensis 

100. Ovary more than 1 mm high. 

101. Seeds straight 42. S. ophirensis 

101. Seeds not straight. 
102. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 

102. Seed and embryo different 42. S. ophirensis 

94. Ovary hairy. 

103. Leaves shorter than 5 cm. Petiole 3-4 mm 10. S. brachybotrys 

103. Leaves longer than 5 cm. 
104. Calyx glabrous. 
105. Disk glabrous. 
106. Calyx lobes not becoming longer by tearing. Seeds not straight. 

16-4. S. cochinchinensis ssp. leptophylla 

106. Calyx lobes becoming longer by tearing. Seeds straight 20. S. crassipes 

105. Disk hairy. 

107. Seeds not straight. Bracts and bracteoles persistent. 

16-4. S. cochinchinensis ssp. leptophylla 
107. Seeds straight. Bracts and bracteoles caducous. 

108. Corolla 5-6 mm 21. S. cylindracea 

108. Corolla 8-10 mm 32. S. junghuhnii 

104. Calyx hairy. 
109. Bracts caducous. 
110. Calyx longer than 1 mm. Style base hciry. 
111. Petiole 3-4 mm. Inflorescence a (basally branched) spike. Ovary c. 1 mm high. Disk 
inconspicuous. Fruits c. 10 mm long, 1 -celled. Seeds not straight . 10. S. brachybotrys 
111. Petiole more than 5 mm. Inflorescence a panicle. Ovary 1-1 ' , mm high. Disk clearly 

present. Fruits 15 mm long, 3-celled. Seeds straight 21. S. cylindracea 

110. Calyx to c. 1 mm long. Style base glabrous 42. S. ophirensis 

109. Bracts persistent. 

1 1 2. Petiole 0-5 mm 42. S. ophirensis 

1 12. Petiole more than 5 mm. 
1 13. Leaves crowded towards the end of the twigs, the latter tapering off, at least 5 mm 

beneath the leaves 44. S. polyandra 

1 13. These characters not combined. 
114. Nerves 13-20 pairs. Intramarginal vein absent. Leaves 21-62 cm 26. S. gigantifolia 
I 14. Nerves 4-13 pairs. I eaves 4-23 cm. 
1 15. Disk glabrous. 
116. Mr acts longer than I mm 16 4. S. cochinchinensis tap. leptophylla 

1 16. Bracti to c. I nam long 

I 17. C alyx lobes to c. ' ., mm long. Stone with low ridges 24. S. lilipes 

117. Calyx lobes longer than ' .. mm StOOC with high lengthwise Interrupted iule.es. 

42. S. ophirensis 
115. Disk hairy. 
I IK. Nerves 4 7 puns, fruits ampulliform with long beak. r. 7 by 5 mm 

42. S. ophirensis 
118. These character! not combined . 16 4. S. cochinchinensis up, leptophylla 



222 Flora Malesiana [ser. I, vol. 8 2 

93. Calyx and ovary glabrous. 

119. Inflorescence terminal 46. S. pyriflora 

119. Inflorescence axillary. 
120. Bracts caducous. 
121. Nerves more than 10 pairs. 
122. Inflorescence a (basally branched) spike, forming a cone in bud. Fruits more than 20 mm 

long 19. S. costata 

122. Inflorescence not a spike. Fruits less than 20 mm long. 

123. Bracts and bracteoles glabrous 46. S. pyriflora 

123. Bracts and bracteoles hairy. 

124. Leaf margin entire. Disk inconspicuous 36. S. maliliensis 

124. Leaf margin not entire. Disk clearly present. 
125. Bracts shorter than 3 mm. Stamens less than 100. Corolla c. 4 mm 

11. S. brandisii 
125. Bracts longer than 3 mm. Stamens more than 100. Corolla c. 5 mm long. 

lib. S. brandisii var. pseudoclethra 
121. Nerves less than 10 pairs. 
126. Disk hairy. 

127. Stamens 15-40. Petiole 1-3 mm 34. S. lancifolia 

127. Stamens more than 40. Petiole more than 5 mm. 
128. Ovary to c. 1 mm high. 
129. Inflorescence a (basally branched) lax spike. Bracts to c. 1 mm long 

25. S. gambliana 
129. Inflorescence not a spike. Bracts longer than 1 mm. 
130. Inflorescence a (basally branched) raceme. Stamens 40-c. 60. Calyx lobes becoming 

longer by tearing. Style base glabrous 14. S. celastrifolia 

130. Inflorescence a panicle of racemes. Stamens more than 100. Calyx lobes not becom- 
ing longer by tearing. Style base hairy 39. S. nivea 

128. Ovary more than 1 mm high. 

131. Petiole 3-4 mm 9. S. borneensis 

131. Petiole more than 5 mm. 
132. Terminal buds glabrous. 
133. Inflorescence a (basally branched) raceme. Calyx IV2 rnm long. Style base glabrous. 

Fruits c. 10 mm long 8. S. batakensis 

133. Inflorescence a fascicle or very short spike. Calyx c. 2 mm long. Style base hairy. 

Fruits 12-16 mm long 53. S. tricoccata 

132. Terminal buds hairy. Inflorescence a panicle. Calyx longer than 2 mm, lobes longer 
than IV2 rnm. Style base hairy. Fruits more than 10 mm long . 21. S. cylindracea 
126. Disk glabrous. 
134. Fruits 2-5-celled. 
135. Inflorescence a fascicle or a very short spike. Ovary more than 1 mm high. 

53. S. tricoccata 
135. Inflorescence a (basally branched) raceme. Ovary to c. 1 mm high. 
136. Inflorescence axis hairy. Corolla more than 4 mm long. Calyx lobes becoming 

longer by tearing. Stone smooth. Seeds not straight 14. S. celastrifolia 

136. Inflorescence axis glabrous. Corolla shorter than c. 4 mm. Calyx lobes not becoming 
longer by tearing. Stone with ridges or grooves. Seeds straight 27. S. glabriramifera 
134. Fruits 1 -celled. 

137. Reticulation fine. Ovary 2-3 mm high 12. S. buxifolia 

137. Reticulation coarse. Ovary 1-2 mm high. 

138. Inflorescence much branched 16-3. S. cochinchinensis ssp. thwaitesii 

138. Inflorescence simple 51. S. sumuntia 

120. Bracts persistent. 
139. Leaves shorter than 5 cm. 
140. Inflorescence only 1 -flowered. Bracts several. 

141. Bracts shorter than 3 mm. Corolla shorter than c. 4 mm. Ovary 1-2 mm high. Stamens 
less than 30. Stone smooth. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 

141. Bracts longer than 3 mm. Corolla more than 4 mm long. Ovary more than 2 mm high. 
Stamens more than 50. Stone with ridges or grooves. Seed and embryo not uncinately 
curved towards the base 12. S. buxifolia 

140. Inflorescence more-flowered. Bract 1. 

142. Petiole 0-5 mm. 

143. Corolla 5-7 mm 57. S. wbitfordii 

143. Corolla shorter. 
144. Leaf index less than 2. Acumen shorter than 5 mm. Bracts longer than 1 mm. 

16. S. cochinchinensis 

144. Leaf index more than 2. Acumen longer than 5 mm 34. S. lancifolia 

142. Petiole more than 5 mm. 



1977] Symplocaceae (Nooteboom) 223 

145. Inflorescence a basally branched raceme. Corolla 5-7 mm long . . 57. S. whitfordii 
145. Inflorescence a (basally branched) spike. Corolla shorter than c. 4 mm. 

16-4. S. cochinchinensis ssp. ieptophylla 
139. Leaves longer than 5 cm. 

146. Petiole 0-5 mm 34. S. lancifolia 

146. Petiole more than 5 mm. 
147. Inflorescence not a spike. 
148. Inflorescence not a fascicle. 
149. Reticulation fine. Ovary c. 1 mm high 18. S. composiracemosa 

149. Reticulation coarse. Ovary more than 1 mm high 

16-3. S. cochinchinensis ssp. thwaitesii 
148. Inflorescence a fascicle. 

150. Disk glabrous. 

151. Petiole 12-17 mm. Ovary more than 1 mm high. Nerves 8-12 pairs. Fruits 10-12 mm 
long 5. S. atjehensis 

151. Petiole 5-12 mm. Ovary c. 1 mm high. Nerves 10-16 pairs. Fruits 7-10 mm long. 

28. S. glomerata 
150. Disk hairy. 

152. Leaves obovate, longer than 10 cm. Acumen longer than 5 mm. Inflorescence axis 
hairy. Calyx regularly 5-lobed. Fruits ovoid or obovoid, 1 -celled. Seeds 1, not 
straight 16-4. S. cochinchinensis ssp. Ieptophylla 

152. Leaves elliptic or circular, shorter than c. 10 cm. Acumen shorter than 5 mm. In- 
florescence axis glabrous. Calyx 2-4-lobed or symmetrically cleft. Fruits cylindric or 

ellipsoid, 2-5-celled. Seeds more than 1, straight 40. S. obovatifolia 

147. Inflorescence a (basally branched) spike. 
153. Twigs (exceptionally) thick. 
154. Terminal buds hairy. Disk hairy .... 16-4. S. cochinchinensis ssp. Ieptophylla 
154. Terminal buds glabrous. Disk glabrous. 
155. Inflorescence axis hairy. Bracts 2-3 mm, hairy. Bracteoles hairy. Calyx 172-2 mm 

long. Fruits c. 10 mm long 6. S. barisanica 

155. Inflorescence axis glabrous. Bracts 5-7 mm, glabrous. Bracteoles glabrous. Calyx 

longer than 2 mm. Fruits c. 13 mm long 45. S. pulvinata 

153. Twigs not (exceptionally) thick. 
156. Calyx 2-4-lobed or symmetrically cleft. 
157. Petiole 15-25 mm 54. S. trisepala 

157. Petiole 7-12 mm 40. S. obovatifolia 

156. Calyx regularly 5-lobed. 

158. Base angle to 20-30°. Leaf index 3'/ 2 -5. Nerves 11-13 pairs . . 29. S. goodeniacea 
158. Base angle more than 30°. Leaf index less than V/ 2 . Nerves at most 11 pairs. 

159. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. Ieptophylla 
1 59. Seed and embryo twice curved 16-2. S. cochinchinensis ssp. laurina 



KEYS TO FLOWERING MATERIAL ARRANGED BY ISLANDS AND ISLAND GROUPS 

Sumatra 

1. Midrib prominent on the upper surface. 
2. Twigs hairy 4. S. anomala 

2. Twigs glabrous 35. S. lucida 

I. Midrib impressed in the upper surface. 

3. Corolla hairy 41. S. odoratissima 

3. Corolla glabrous. 

4. Underside of leaves glabrous. 
5. Twigs hairy. 

6. Leaves distichous. Calyx 2-4-lobed or symmetrically cleft, calyx lobes becoming longer by tearing. 

33. S. laetcviridis 
6. Leaves spirally arranged. 
7. Leaves crowded towards the end of the twigs. Twigs Ihick, tapering towards apex. Petiole more 
than 20 mm. C orolla more than 7 nun long. Apex of leaves rounded or acute . 44. S. polyandra 
7. Leaves evenly distributed. 

8. Calyx and ovary glabrous 5. S. atjehensis 

8. Calyx and or ovary hairy. 
9 I eaves longer than 10 cm. Nerves more than 10 pairs. Inflorescence a (basally branched) spike. 
Hracts persistent. Ovary glabrous, to , I mm Ugh, calyx lobes longer than I'/itntn. Disk 
glabrous, clearly present I nuts ainpullilorm, I -celled. 

16 I. S. cochinchinensis ssp. cochinchinensis 



224 Flora Malesiana [ser. I, vol. 8 2 

9. Leaves shorter than c. 10 cm. Nerves less than 10 pairs. Inflorescence a raceme. Bracts 
caducous. Ovary hairy, 1-2 mm high, calyx lobes V2-IV2 mm long. Disk hairy, inconspicuous. 

Fruits ellipsoid, 3-celled 47. S. robinsonii 

5. Twigs glabrous. 

10. Nerves 4-5 pairs 42. S. ophirensis 

10. Nerves more than 5 pairs. 
11. Calyx and/or ovary hairy. 

12. Leaves distichous 33. S. laeteviridis 

12. Leaves spirally arranged. 

13. Leaves crowded towards the end of the tapering-off twigs 44. S. polyandra 

13. Plant different 42. S. ophirensis 

11. Calyx and ovary glabrous. 
14. Inflorescence not a spike. 

15. Inflorescence a fascicle. Stamens c. 50 5. S. atjehensis 

15. Inflorescence a raceme. 

16. Stamens 40-60 14. S. celastrifolia 

16. Stamens c. 100 8. S. batakensis 

14. Inflorescence a (basally branched) spike. 

17. Twigs thick. Terminal buds large 6. S. barisanica 

17. Twigs not thick. Terminal buds small 16-2. S. cochinchinensis ssp. laurina 

4. Underside of leaves hairy. 
18. Twigs glabrous. 

19. Leaves distichous. Petiole 1-4 mm. Corolla 3-5 mm 33. S. laeteviridis 

19. Leaves spirally arranged. Petiole more than 5 mm. 
20. Inflorescence a fascicle. Bracts persistent, c. 2 mm. Calyx glabrous. Calyx lobes not becoming 

longer by tearing 5. S. atjehensis 

20. Inflorescence a spike forming a cone in bud. Bracts caducous, 3-5 mm. Calyx hairy. Calyx lobes 

becoming longer by tearing 48. S. rubiginosa 

18. Twigs hairy. 
21. Leaves distichous. 
22. Calyx usually hairy. Inflorescence a fascicle. Bracts persistent. Ovary c. 1 mm long. Calyx 
c. 1 mm long. Calyx lobes not becoming longer by tearing. Style base hairy . 23. S. fasciculata 

22. Calyx often glabrous. Inflorescence a raceme or panicle of racemes. Bracts caducous. Ovary 
I-IV2 mm high. Calyx 2-3 mm. Calyx lobes becoming longer by tearing. Style base glabrous 

33. S. laeteviridis 
21. Leaves spirally arranged. 

23. Calyx and ovary glabrous. 

24. Leaf index 2 to 3. Nerves 8-12 pairs 5. S. atjehensis 

24. Leaf index more than 3. Nerves 6-9 pairs. 

25. Nerves less than 10 pairs 15. S. cerasifolia 

25. Nerves more than 10 pairs 15b. S. cerasifolia var. grandifolia 

23. Calyx and/or ovary hairy. 
26. Upper side of leaves hairy (pulverulent). Leaf margin and petiole beset with many closely 

spaced vesicular glands 3. S. adenophylla 

26. Upper side of leaves glabrous. Leaf margin and petiole different. 
27. Ovary glabrous. 
28. Nerves less than 10 pairs. Bracts caducous. Ovary I-IV2 mm high. Calyx 2V 2 -4 mm. Calyx 
lobes becoming longer by tearing 15. S. cerasifolia 

28. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm long. Calyx 1 to 2 mm long. 
Calyx lobes not becoming longer by tearing. 

16-1. S. cochinchinensis ssp. cochinchinensis 
27. Ovary hairy. 

29. Leaves longer than 15 cm. 

30. Nerves 10-13 pairs 15b. S. cerasifolia var. grandifolia 

30. Nerves 12-17 pairs 48. S. rubiginosa 

29. Leaves shorter than c. 1 5 cm. 

31. Bracts to c. 1 mm long. 

32. Leaf margin (and petiole) beset with closely spaced glands 3. S. adenophylla 

32. Leaf margin (and petiole) often glandular but glands not closely spaced. 
33. Nerves less than 10 pairs. Reticulation coarse. Inflorescence a fascicle. Bracts persistent. 

Ovary c. 1 mm high. Style base hairy 23. S. fasciculata 

33. Nerves more than 10 pairs. Reticulation fine. Inflorescence a (basally branched) raceme. 
Bracts caducous. Ovary more than 1 mm high. Style base glabrous 47. S. robinsonii 
31. Bracts longer than 1 mm. 

34. Angle of leaf base c. 90° 50. S. sumatrana 

34. Angle of leaf base less than 60° 47. S. robinsonii 



1977] Symplocaceae (Nooteboom) 225 

Malay Peninsula 

1. Midrib prominent on the upper surface. 
2. Twigs hairy. 
3. Leaves evenly distributed, underside glabrous 4. S. anomala 

3. Leaves crowded towards the end of the twigs, minutely sparsely appressedly hairy beneath. 

37. S. wikstroemifolia 
2. Twigs glabrous. 

4. Leaves crowded towards the end of the twigs 37. S. wikstroemifolia 

4. Leaves evenly distributed 35. S. lucida 

1. Midrib impressed on the upper surface. 

5. Corolla hairy 41. S. odoratissima 

5. Corolla glabrous. 
6. Twigs hairy. 
7. Leaves distichous. 

8. Underside of leaves glabrous 33. S. laeteviridis 

8. Underside of leaves hairy. 
9. Bracts persistent. 
10. Inflorescence a true fascicle. Ovary c. 1 mm high. Calyx 1 mm long ... 23. S. fasciculata 
10. Inflorescence a short, often clustered spike. Ovary 1-2 mm high. Calyx 1V2-4 mm long 

20. S. crassipes 

9. Bracts caducous 33. S. laeteviridis 

7. Leaves spirally arranged. 
11. Upper side of leaves hairy. 
12. Angle of leaf base more than 90°. Bracts and bracteoles caducous. Hairs on twigs more than 

2 mm long 13. S. calycodactylos 

12. Angle of leaf base less than 90°. 
13. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 mm long. Calyx to 
c. 1 mm long, calyx lobes 1 / 2 -l mm long. Disk glabrous. Style base not conical. Fruits to c. 

10 mm long 3. S. adenophylla 

13. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than 
1 mm. Calyx l'/j-4mm, calyx lobes longer than l'/ 2 mm. Disk hairy. Style base conical. 

Fruits 13-18 mm long 20. S. crassipes 

11. Upper side of leaves glabrous. 

14. Leaves crowded towards the end of the twigs 37. S. wikstroemifolia 

14. Leaves evenly distributed. 
15. Underside of leaves glabrous. 
16. Calyx glabrous. Inflorescence a fascicle. Disk pulvinate or cylindric. Fruits cylindrical. Seeds 
straight 28. S. glomerata 

16. Calyx hairy. Inflorescence a (basally branched) spike. Disk annular. Fruits ampulliform. 
Seeds not straight 16-1. S. cochinchinensis ssp. cochinchinensis 

15. Underside of leaves hairy (pulverulent, nearly glabrous, in S. adenophylla). 

17. Calyx and ovary glabrous. 

18. Inflorescence a spike 15. S. cerasifolia 

18. Inflorescence a fascicle 28. S. glomerata 

17. Calyx and or ovary hairy. 

19. Ovary glabrous. 

20. Nerves less than 10 pairs. Bracts and bracteoles caducous. Ovary 1-1 V2 mm high, calyx 
longer than 2 mm, calyx lobes 1-1 l /a mm long, becoming longer by tearing. Fruits 22-40 
mm long, 3-celled 15. S. cerasifolia 

20. Nerves more than 10 pairs. Bracts and bracteoles persistent. Ovary to c. 1 mm high, calyx 
c. 2 mm long, calyx lobes longer than 1' 2 mm, not becoming longer by tearing. Fruits 
5-7 mm long, 1 -celled 16-1. S. cochinchinensis ssp. cochinchinensis 

19. Ovary hairy. 

21. Ovary more than 1 mm high. 

22. Leaf margin (and petiole) beset with closely spaced glands. Bracts to c. 1 mm long. Calyx 

1 mm long 3. S. adenophylla 

22. Leaf margin (and petiole) often glandular but glands not closely spaced. Bracts longer than 

I mm. 

23. Nerves (, II pairs. Reticulation faintly prominent. Bracts and bracteoles persistent. 

( alyx lobes not becoming longer by tearing. I nuts 13 IN mm long . 20. S. crassipes 

23. Nerves 12 17 pairs. Reticulation much prominent. Bracts and bracteoles caducous. 

( alyx lobes becoming longer by tearing. Fruit! to c. 10 mm long . . 48. S. raUgtaOM 

21. Ovary to c. I mm high. 

24 I Qgef than 15 cm 20. S. crassipes 

24. leaves shorter than C. IScm. 

25. (Reticulation line.) Brads and hi. icteoles caducous 42. S. ophirensis 

25. (Reticulation coarse.) Bracts persistent. 



226 Flora Malesiana [ser. I, vol. 8 2 

26. Inflorescence an often clustered short spike. Bracts 1-4 mm. Calyx 2Vz-3 mm. Stamens 

more than 60. Fruits not ampulliform, 13-18 mm long 20. S. crassipes 

26. Inflorescence a fascicle. Bracts to c. 1 mm long. Calyx c. 1 mm long. Stamens 12-35. 

Fruits ampulliform, 5-7 mm long 23. S. fasciculata 

6. Twigs glabrous. 
27. Nerves more than 10 pairs. 
28. Underside of leaves hairy. 
29. Inflorescence a spike 48. S. rubiginosa 

29. Inflorescence a fascicle 28. S. glomerata 

28. Underside of leaves glabrous. 

30. Petiole more than 20 mm. Inflorescence a spike or a cone. Fruits 3-celled 

7. S. barringtoniifolia 
30. Petiole less than 20 mm. 

31. Inflorescence terminal 46. S. pyriflora 

31. Inflorescence axillary. 

32. Calyx and/or ovary hairy. Corolla 2-5 mm 42. S. ophirensis 

32. Calyx and ovary glabrous. 

33. Intramarginal vein far from margin. Inflorescence a fascicle. Bracts persistent, hairy, shorter 
than 3 mm. Ovary c. 1 mm high, calyx 1-2 mm long, calyx lobes not becoming longer by 
tearing. Corolla 4-5 mm. Stamens less than 50. Fruits 7-10 mm long. . 28. S. glomerata 
33. Intramarginal vein close to margin. Inflorescence a raceme or panicle of racemes. Bracts 
caducous, glabrous, longer than 3 mm. Ovary IV2-2 mm high, calyx 3-5 mm, calyx lobes 
becoming longer by tearing. Corolla 8-10 mm long. Stamens c. 100 or more. Fruits c. 15 mm 

long 46. S. pyriflora 

27. Nerves less than 10 pairs. 
34. Underside of leaves hairy. 

35. Leaves crowded towards the end of the twigs 37. S. wikstroemifolia 

35. Leaves evenly distributed. 

36. Bracts persistent. Disk glabrous. Fruits 13-18 mm long 20. S. crassipes 

36. Bracts caducous. Disk hairy. Fruits 7-12 mm long 33. S. laeteviridis 

34. Underside of leaves glabrous. 
37. Calyx and/or ovary hairy. 
38. Disk hairy. 
39. Leaves distichous 33. S. laeteviridis 

39. Leaves spirally arranged 42. S. ophirensis 

38. Disk glabrous. 

40. Inflorescence a short, often clustered spike. Bracts persistent 20. S. crassipes 

40. Inflorescence a raceme. Bracts caducous 51. S. sumuntia 

37. Calyx and ovary glabrous. 
41. Inflorescence a (basally branched) spike. Bracts persistent 

16-2. S. cochinchinensis ssp. laurina 
41. Inflorescence not a spike. Bracts caducous. 

42. Inflorescence a panicle of racemes. Stamens more than 100 39. S. nivea 

42. Inflorescence a (basally branched) raceme. Stamens 25-60. 

43. Calyx lobes becoming longer by tearing 14. S. celastrifolia 

43. Calyx lobes not becoming longer by tearing 51. S. sumuntia 

Java & The Lesser Sunda Islands 

1. Corolla hairy 41. S. odoratissima 

1. Corolla glabrous. 

2. Midrib prominent on the upper surface 35. S. lucida 

2. Midrib impressed in the upper surface. 
3. Twigs hairy. 

4. Leaves distichous 23. S. fasciculata 

4. Leaves spirally arranged. 
5. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 

3. S. adenophylla.) 16-1. S. cochinchinensis ssp. cochinchinensis 

5. Underside of leaves hairy. 

6. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

6. Upper side of leaves glabrous. 
7. Ovary glabrous, calyx longer than 1 mm. Bracts longer than 1 mm 

16-1. S. cochinchinensis ssp. cochinchinensis 
7. Ovary hairy. 
8. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Ovary 

1-2 mm high 3. S. adenophylla 

8. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. 
Ovary c. 1 mm high 23. S. fasciculata 



1977] Symplocaceae (Nooteboom) 227 

3. Twigs glabrous. 
9. Calyx and or ovary hairy. 
10. Ovary hairy. Inflorescence a raceme. Calyx glabrous 32. S. junghuhnii 

10. Ovary glabrous. Inflorescence a spike. Calyx hairy. 

16-1. S. cochinchinensis ssp. cochinchinensis 
9. Calyx and ovary glabrous. 

1 1 . Inflorescence axis glabrous. 

12. Petiole less than 20 mm 16-2. S. cochinchinensis ssp. laurina 

12. Petiole more than 20 mm 16-4. S. cochinchinensis ssp. leptophy Ha 

1 1 . Inflorescence axis hairy. 

13. Nerves less than 10 pairs 16. S. cochinchinensis 

13. Nerves more than 10 pairs. 

14. Angle of leaf base 25-40°. Inflorescence a (basally branched) spike, forming a cone in bud. 

19. S. costata 
14. Angle of leaf base more than 60°. Inflorescence a raceme 11. S. brandisii 

Borneo 

1. Corolla hairy 41. S. odoratissima 

1. Corolla glabrous. 

2. Midrib flat or prominent on the upper surface 4. S. anomala 

2. Midrib impressed in the upper surface. 
3. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 

3. 5. adenophylla). 
4. Twigs hairy. 

5. Leaves distichous 33. S. laeteviridis 

5. Leaves spirally arranged. 

6. Leaves 4-6 cm. Petiole 3-4 mm 10. S. brachybotrys 

6. Leaves longer than 6 cm. Petiole more than 5 mm. 
7. Leaves evenly distributed. Twigs not thick, cylindrical. Leaf margin not entire. Petiole less than 
20 mm. Ovary glabrous, to c. 1 mm high, calyx 1-2 mm long. Corolla 3-5 mm. Fruits ampulli- 
form, 1 -celled. Seed 1, not straight. Apex of leaves acuminate. 

16-1. S. cochinchinensis ssp. cochinchinensis 

7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Leaf margin 

entire. Petiole more than 20 mm. Ovary hairy, c. 2 mm high, calyx 2-3 mm long. Corolla 

8-10 mm long. Fruits ellipsoid, 3-celled. Seeds more than 1, straight. Apex of leaves rounded or 

acute 44. S. polyandra 

4. Twigs glabrous. 
8. Calyx and/or ovary hairy. 

9. Leaves distichous. Calyx glabrous 33. S. laeteviridis 

9. Leaves spirally arranged. 
10. Inflorescence a 1-3-flowered spike. Bracts caducous. Stamens c. 100. Petiole 3-4 mm. Leaves 

4-6 cm 10. S. brachybotrys 

10. These characters not combined. 

11. Leaves 15-50 mm long 12. S. buxifolia 

1 1 . Leaves longer than 5 cm. 
12. Petiole more than 20 mm. Twigs thick. Inflorescence a spike. Calyx 3— 3Vi mm 

7. S. barringtoniifolia 
12. Petiole less than 20 mm. Inflorescence a raceme or a spike. Calyx to c. 1 mm long 

42. S. ophirensis 
8. Calyx and ovary glabrous. 

13. Nerves more than 10 pairs 29. S. goodeniacea 

13. Nerves less than 10 pairs. 
14. Bracts persistent. 
15. Leaves shorter than 5 cm 12. S. buxifolia 

15. Leaves longer than 5 cm 16-2. S. cochinchinensis ssp. laurina 

14. Bracts caducous. 

16. Ovary to c. I mm high. Apex of leaf mostly abruptly acuminate. 

17. Leaf margin entire. Inflorescence axis glabrous. Petiole 5 10 mm . . . 25. S. gambliana 

17. Leaf margin not entire. Inflorescence axis hairy. Petiole 3-15 mm 14. S. celastrifolia 
16. Ovary more than I mm high I eal apex usually not or faintly acuminate. 

I - leaf margin entire. Calyx *J 4 I mm long. Disk hairy. Petiole 3 4 mm . 9. S. borncensis 

18. Leaf margin QOt entire ( alyx lunger than I mm. Disk glabrous or the Style base hairy. 
19. Leaves shorter than 3 Cfll ami (U iinien shorter than 5 mm. Reticulation fine. Inflorescence a 

few-flowered raceme. Bract! lunger than 3 mm. Ovary 2-3 mm high, calyx 2 5 mm long. 

Style bate glabrous. I nuts I -celled 12. S. buxifolia 

19. Leaves longer than 5 cm and acumen longer than 5 mm. Reticulation coarse. Inflorescence 

i i ' J nun ( rvary c. 2 mm high, calyx e. 2 mm long. 
Style base hairy, fruits Vcellecl. Stone smooth. Seeds more than I . . 53. S. tricoccata 



228 Flora Malesiana [ser. I, vol. 8 2 

3. Underside of leaves hairy. 
20. Leaves distichous. 

21. Calyx symmetrically teared when older 33. S. laeteviridis 

21. Calyx regular. 
22. Stamens 25-35. Inflorescence a true fascicle. Petiole 2-8 mm. Leaves 5-18 cm. Ovary c. 1 mm 

high. Calyx c. 1 mm long 23. S. fasciculata 

22. Stamens more than 40. Flowers solitary or in a raceme or panicle. Leaves 2-9 cm. Calyx 1-3 mm. 
23. Nerves 7-11 pairs. Petiole 3-4 mm. Leaves 4-9 cm. Stamens more than 90. Calyx l 3 / 4 -3 mm 

17. S. colombonensis 
23. Nerves 3-8 pairs. Stamens 40-100, but when more than 90 petiole 1-2 mm. 

24. Petiole 2-4 mm. Leaves 2-3V 2 cm. Calyx c. 3 mm 52. S. trichomarginalis 

24. Petiole 1-2 mm. Leaves 2*/ 2 -7 cm. Calyx 1-2 mm. 

25. Calyx c. 2 mm long 58. S. zizyphoides 

25. Calyx I-IV2 mm long. 

26. Ovary 1 mm high 31. S. johniana 

26. Ovary l'/^mmhigh 22. S. deflexa 

20. Leaves spirally arranged. 
27. Twigs glabrous. 

28. Leaves 4-6 cm 10. S. brachybotrys 

28. Leaves longer than 6 cm. 

29. Leaves 7-16 cm. Nerves 6-9 pairs 15. S. cerasifolia 

29. Leaves 15-45 cm. Nerves 12-17 pairs 48. S. rubiginosa 

27. Twigs hairy. 

30. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

30. Upper side of leaves glabrous. 

31. Calyx and ovary glabrous 15. S. cerasifolia 

31. Calyx and/or ovary hairy. 
32. Ovary glabrous. 

33. Nerves less than 10 pairs. Bracts caducous. Ovary 1-1 V2 mm high, calyx longer than 2 mm. 
Fruits ellipsoid, 22-40 mm long, 3-celled 15. S. cerasifolia 

33. Nerves more than 10 pairs. Bracts persistent. Ovary to c. 1 mm high, calyx 1-2 mm long. 
Fruits ampulliform, 5-7 mm long, 1 -celled. 

16-1. S. cochinchinensis ssp. cochinchinensis 
32. Ovary hairy. 

34. Leaves longer than 15 cm 48. S. rubiginosa 

34. Leaves shorter than c. 15 cm. 

35. Inflorescence only 1 -flowered. 

36. Angle of leaf base more than 90° 31. S. johniana 

36. Angle of leaf base less than 90°. 

37. Bracts 1. Calyx 1-2 mm long 20. S. crassipes 

37. Bracts several. Calyx c. 3 mm 10. S. brachybotrys 

35. Inflorescence more-flowered. 

38. Bracts caducous. Petiole 3-4 mm. Nerves 6-9. Stamens c. 100 . . 10. S. brachybotrys 
38. Bracts persistent. Petiole 2-12 mm. Nerves 3-12. Stamens 12-more than 100. 
39. Calyx to c. 1 mm long. Stamens 12-50. 
40. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. 

Ovary 1-2 mm high 3. S. adenophylla 

40. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. 

Ovary c. 1 mm high 23. S. fasciculata 

39. Calyx longer than 1 mm. Stamens 25-100 20. S. crassipes 

Philippines 

1 . Leaves verticillate 55. S. verticillifolia 

1. Leaves not verticillate. 
2. Midrib prominent in the upper surface. 
3. Twigs glabrous. Petiole more than 5 mm 35. S. lucida 

3. Twigs hairy. Petiole 1-5 mm 34. S. lancifolia 

2. Midrib impressed on the upper surface. 

4. Corolla hairy (in the Philippines sometimes nearly glabrous!) 41. S. odoratissima 

4. Corolla glabrous. 

5. Twigs hairy. 

6. Leaves distichous 23. S. fasciculata 

6. Leaves spirally arranged. 

7. Calyx divided into three 2*/ 2 mm long lobes 54. S. trisepala 

7. Calyx not so. 
8. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards the apex. Fruits 
3-celled. Apex of leaves rounded or acute 44. S. polyandra 



1977] Symplocaceae (Nooteboom) 229 

8. Leaves evenly distributed. 
9. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular 
glands: 3. 5. adenophylla). 
10. Ovary hairy, c. 1 ' 2 mm high. Inflorescence a lax raceme. Bracts to c. 1 mm long. Calyx lobes 
c. ' 2 mm long. Stamens c. 25 24. S. filipes 

10. Ovary glabrous (hidden between bracts!). Inflorescence a spike. 

16-1. S. cochinchinensis ssp. cochinchinensis 
9. Underside of leaves hairy. 

11. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

11. Upper side of leaves glabrous. 

12. Ovary glabrous. 
13. Leaves longer than 10 cm. Petiole more than 5 mm. Nerves more than 10 pairs. Calyx 
lobes longer than l 1 ; 2 mm. Stamens more than 30. Disk glabrous. Style base glabrous. 
Fruits ampulliform 16-1. S. cochinchinensis ssp. cochinchinensis 

13. Leaves shorter than c. 10 cm. Petiole 1-5 mm. Nerves 4-11 pairs. Calyx lobes 'j-l 1 ^ mm 
long. Stamens 15-40. Disk hairy. Style base hairy. Fruits not ampulliform 

34. S. lancifolia 
12. Ovary hairy. 

14. Leaf margin (and petiole) beset with closely spaced glands .... 3. S. adenophylla 
14. Leaf margin (and petiole) often glandular but glands not closely spaced. 

15. Style base glabrous. 
16. Bracts to c. 1 mm long. Calyx lobes c. ' 2 mm long, not triangular . . 24. S. filipes 

16. Bracts 2-3 mm. Calyx lobes 1-1 l 2 mm, triangular 56. S. vidalii 

15. Style base hairy. 

17. Intramarginal vein present. Inflorescence a fascicle. Fruits ampulliform 

23. S. fasciculata 
17. Intramarginal vein absent. Inflorescence a (basally branched) spike. Fruits ellipsoid to 

orbicular 34. S. lancifolia 

5. Twigs glabrous. 
18. Calyx and ovary glabrous. 
19. Inflorescence a (basally branched) spike. 

20. Acumen longer than 5 mm 34. S. lancifolia 

20. Acumen shorter than 5 mm. 
21. Angle of leaf base less than 60°. Bracts 2-3 mm. Calyx lobes c. 2 mm long. Style base hairy. 

40. S. obovatifolia 
21. Angl: of leaf base c. 90°. Bracts 3-5 mm. Calyx lobes longer than 2 l / 2 mm. Disk glabrous. 

54. S. trisepala 
19. Inflorescence not a spike. 
22. Bracts and bracteoles persistent. 
23. Leaves 2-5 3 4 cm. Inflorescence a (basally branched) raceme. Bracts longer than 3 mm. Calyx 
regularly 5-lobed, calyx lobes semi-ovate. Fruits ovoid, 5-7 mm long, 1 -celled. 

57. S. whitfordii 

23. Leaves 7 1 2 -ll cm. Inflorescence a fascicle. Bracts 2-3 mm long. Calyx 3-lobed, the lobes 
semi-elliptic. Fruits ellipsoid, 1 1 mm long, 3-celled 40. S. obovatifolia 

22. Bracts and bracteoles caducous. 

24. Inflorescence axis glabrous. Corolla 3^* mm. Calyx lobes not becoming longer by tearing. 

27. S. glabriramifera 

24. Inflorescence axis hairy. Corolla 4-6 mm 14. S. celastrifolia 

18. Calyx and or ovary hairy. 
25. Inflorescence a spike. Ovary glabrous. 
26. Ovary ' 2 -l mm high 16-1. S. cochinchinensis ssp. cochinchinensis 

26. Ovary 2 1 2 mm high 42. S. ophirensis 

25. Inflorescence a (sometimes compound) raceme. Ovary hairy. 

27. Inflorescence a very lax raceme of 4-10 cm. Pedicels slender, 2-15 mm. Axis of raceme sparsely 
pulverulent-puberulous. Stamens c. 25 24. S. filipes 

27. These characters not combined 42. S. ophirensis 

Celebes & The Moluccas 

I. Corolla hairy 41. S. odoratissima 

1. Corolla glabrous. 

2. Midrib prominent on the upper surface 35. S. lucida 

2. Midrib impressed in the upper surface. 
3. Twigs hairy. 

4. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 
3. .V. adenophylla). 

5. Leaves distichous. Mracts caducous 33. S. laetcviridis 

5. Leaves spirally arranged. 



230 Flora Malesiana [ser. I, vol. 8 2 

6. Calyx and ovary glabrous 16-4. S. cochinchinensis ssp. leptophylla 

6. Calyx and/or ovary hairy. 
7. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. 

44. S. polyandra 
7. Leaves evenly distributed. 

8. Ovary hairy 16-4. S. cochinchinensis ssp. leptophylla 

8. Ovary glabrous. 

9. Ovary to c. 1 mm high 16-1. S. cochinchinensis ssp. cochinchinensis 

9. Ovary more than 1 mm high 16-4. S. cochinchinensis ssp. leptophylla 

4. Underside of leaves hairy. 
10. Leaves distichous. 

11. Inflorescence a fascicle. Bracts persistent. Ovary c. 1 mm high, calyx c. 1 mm long, calyx lobes 
not becoming longer by tearing. Style base hairy. Fruits ampulliform . . 23. S. fasciculata 

11. Inflorescence not a fascicle. Bracts caducous. Ovary I-lVa mm high, calyx 2-3 mm, calyx lobes 
becoming longer by tearing. Style base glabrous. Fruits not ampulliform . . 33. S. laeteviridis 

10. Leaves spirally arranged. 

12. Calyx and ovary glabrous 16-4. S. cochinchinensis ssp. leptophylla 

12. Calyx and/or ovary hairy. 

13. Upper side of leaves hairy 3. S. adenophylla 

13. Upper side of leaves glabrous. 

14. Ovary glabrous 16-4. S. cochinchinensis ssp. leptophylla 

14. Ovary hairy. 

15. Calyx glabrous 16-4. S. cochinchinensis ssp. leptophylla 

15. Calyx hairy. 
16. Bracts to c. 1 mm long. 
17. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Ovary 
1-2 mm high. Stone not ampulliform 3. S. adenophylla 

17. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. 
Ovary c. 2 mm high. Stone ampulliform 23. S. fasciculata 

16. Bracts longer than 1 mm. 

18. Bracts caducous. Ovary c. 1 mm high .... 42-lb. S. ophirensis var. densireticulata 
18. Bracts persistent. Ovary more than 1 mm high 16-4. S. cochinchinensis ssp. leptophylla 

3. Twigs glabrous. 
19. Underside of leaves hairy. 
20. Leaves distichous. Petiole 0-5 mm. Bracts and bracteoles caducous . . . . 33. S. laeteviridis 

20. Leaves spirally arranged 16-4. S. cochinchinensis ssp. leptophylla 

19. Underside of leaves glabrous. 

21. Calyx and ovary glabrous. 
22. Ovary to c. 1 mm high. 

23. Inflorescence a raceme. Bracts caducous 14. S. celastrifolia 

23. Inflorescence a (basally branched) spike. Bracts persistent 

16-2. S. cochinchinensis ssp. laurina 
22. Ovary more than 1 mm high. 

24. Twigs thick 16-4. S. cochinchinensis ssp. leptophylla 

24. Twigs not thick. 

25. Inflorescence a raceme. Bracts caducous 36. S. maliliensis 

25. Inflorescence a (basally branched) spike. Bracts persistent. 

26. Disk hairy 16-4. S. cochinchinensis ssp. leptophylla 

26. Disk glabrous 16-2. S. cochinchinensis ssp. laurina 

21. Calyx and/or ovary hairy. 

27. Leaves distichous 33. S. laeteviridis 

27. Leaves spirally arranged. 
28. Ovary glabrous. 
29. Ovary to c. 1 mm high 16-1. S. cochinchinensis ssp. cochinchinensis 

29. Ovary more than 1 mm. high 16-4. S. cochinchinensis ssp. leptophylla 

28. Ovary hairy. 

30. Calyx glabrous 16-4. S. cochinchinensis ssp. leptophylla 

30. Calyx hairy. 

31. Bracts to c. 1 mm long 42. S. ophirensis 

31. Bracts longer than 1 mm 16-4. S. cochinchinensis ssp. leptophylla 

New Guinea 
(inch New Ireland & New Britain) 

1 . Leaves (pseudo-)verticillate 30. S. herzogii 

1. Leaves not verticillate. 
2. Inflorescence only 1 -flowered. 
3. Calyx and ovary glabrous 16-4. S. cochinchinensis ssp. leptophylla 



1977] Symplocaceae (Nooteboom) 231 

3. Calyx and or ovary hairy. 

4. Ovary hairy 49. S. salicioides 

4. Ovary glabrous 38. S. multibracteata 

2. Inflorescence more-flowered. 
5. Calyx and ovary glabrous. 
6. Twigs hairy. 

7. Petiole to 5 mm 16-4. S. cochinchinensis ssp. leptophylla 

7. Petiole more than 5 mm. 

8. Underside of leaves glabrous 16-4. S. cochinchinensis ssp. leptophylla 

8. Underside of leaves hairy. 

9. Upper side of leaves hairy. Ovary c . 3 / 4 mm long 43. S. paucistaminea 

9. Upper side of leaves glabrous. Ovary more than 1 mm high. 

10. Bracts caducous. Calyx lobes becoming longer by tearing 15. S. cerasifolia 

10. Bracts persistent. Calyx lobes not becoming longer by tearing. 

16-4. S. cochinchinensis ssp. leptophylla 
6. Twigs glabrous. 

11. Underside of leaves hairy 1 6 4. S. cochinchinensis ssp. leptophylla 

11. Underside of leaves glabrous. 
12. Bracts caducous. 

13. Bracts longer than 3 mm 16-4. S. cochinchinensis ssp. leptophylla 

13. Bracts shorter than c. 3 mm. 

14. Ovary c. 1 mm long. Inflorescence a (basally branched) raceme .... 14. S. celastrifolia 
14. Ovary more than 1 mm high. Inflorescence different. 

15. Disk hairy. Ovary 1-1 V 2 mm. Inflorescence a panicle 21. S. cylindracea 

1 5. Disk glabrous. Inflorescence an often branched spike. 

16-3. S. cochinchinensis ssp. thwaitesii 
12. Bracts persistent. 
16. Inflorescence not a spike. 
17. Inflorescence a fascicle 16-4. S. cochinchinensis ssp. leptophylla 

17. Inflorescence a panicle 18. S. composiracemosa 

16. Inflorescence a (basally branched) spike. 

18. Twigs not thick. 

19. Leaves 6-12 cm. Petiole 5-25 mm. Inflorescence an (often branched) spike. Flowers bisexual, 
ovary 1-1 V 2 mm, calyx 1 U- i l 4 mm 16 3. S. cochinchinensis ssp. thwaitesii 

19. Plants different. Flowers usually functional unisexual. 

16-4. S. cochinchinensis ssp. leptophylla 
18. Twigs thick. 

20. Terminal buds hairy, small. Acumen longer than 5 mm. Bracts hairy, shorter than 3 mm. 
Disk hairy. Leaves elliptic or circular 16-4. S. cochinchinensis ssp. leptophylla 

20. Terminal buds glabrous, large. Acumen shorter than 5 mm. Bracts glabrous, 5-7 mm. Disk 

glabrous. Leaves obovate 45. S. pulvinata 

5. Calyx and or ovary hairy. 
21. Ovary glabrous. 
22. Ovary hidden by bracts and bracteoles .... 16-1. S. cochinchinensis ssp. cochinchinensis 
22. Ovary not hidden by bracts and bracteoles. 

23. Inflorescence a spike, forming a short cone in bud 15. S. cerasifolia 

23. Inflorescence sometimes a spike, but never forming a cone in bud. 

16-4. S. cochinchinensis ssp. leptophylla 
21. Ovary hairy. 

24. Twigs at least 8 mm thick. Leaves 21-62 cm 26. S. gigantifolia 

24. Twigs thinner. Leaves at most 33 cm, but usually much smaller. 
25. Calyx 2- to 4-lobed or symmetrically cleft. Calyx lobes becoming longer by tearing. 

16-4. S. cochinchinensis ssp. leptophylla 
25. Calyx regularly 5-lobed. 

26. Disk glabrous 16-4. S. cochinchinensis tap. leptophylla 

26. Disk hairy. 

27. Bracts and bracteoles caducous 21. S. cylindracea 

27. Bracts persistent 16 4. S. cochinchinensis ssp. leptophylla 



KEYS TO FRUITINfi MAIIRIAI ARRANdU) II Y ISLANDS AND ISLAND GROUPS 

Sumatra 

I. Midrib prominent on the upper BUlftoC 
2. Twigs hairy. 'Icrnniial bud-, hairy, small. Seeds straight 4. S. anomala 

2. Twigs glabrous. Terminal bud . j'l.ilm hi , \o-d.un\ol 35. S. lucidu 

1. Midrib impressed Oil the upper smi 

3. Underside of leaves glabrous. 



232 Flora Malesiana [ser. I, vol. 8 2 

4. Twigs hairy. 

5. Leaves distichous 33. S. laeteviridis 

5. Leaves spirally arranged. 
6. Leaves crowded towards the end of the twigs. Twigs thick, tapering towards apex. Petiole more 

than 20 mm. Apex of leaves rounded or acute 44. S. polyandra 

6. Leaves evenly distributed. 
7. Fruits ampulliform. Inflorescence a (basally branched) spike. Seeds not straight. 

16-1. S. cochinchinensis ssp. cochinchinensis 
7. Fruits ellipsoid. 

8. Fruits 10-12 mm long. Inflorescence a fascicle. Bracts persistent 5. S. atjehensis 

8. Fruits 7-10 mm long. Inflorescence a (basally branched) raceme. Bracts caducous 

47. S. robinsonii 
4. Twigs glabrous. 

9. Nerves less than 5 pairs 42. S. ophirensis 

9. Nerves more than 5 pairs. 

10. Leaves distichous 33. S. laeteviridis 

10. Leaves spirally arranged. 
11. Inflorescence a (basally branched) spike. 
12. Fruits ovoid to orbicular, 9-10 mm long. Twigs thick. Terminal buds large 6. S. barisanica 

12. Fruits ampulliform to globose, 5-7 mm long .... 16-2. S. cochinchinensis ssp. laurina 
11. Inflorescence not a spike (rarely a cone in bud). 

13. Bracts persistent. 

14. Petiole 12-17 mm. Fruits ellipsoid. Inflorescence a fascicle 5. S. atjehensis 

14. Petiole 3-9 mm. Fruits ampulliform. Inflorescence a raceme or panicle . 42. S. ophirensis 
13. Bracts caducous. 

15. Fruits 1 -celled. Inflorescence a rusty tomentellous panicle 41. S. odoratissima 

15. Fruits 2-3-celled. Inflorescence a raceme. 

16. Leaf margin entire. Fruits c. 10 mm. Seeds straight 8. S. batakensis 

16. Leaf margin not entire. Fruits 4-10 mm, the sterile cells larger than the fertile ones, towards 

the base rilled with air 14. S. celastrifolia 

3. Underside of leaves hairy. 
17. Twigs glabrous. 

18. Leaves distichous. Petiole 1-4 mm 33. S. laeteviridis 

18. Leaves spirally arranged. Petiole more than 10 mm. 

19. Inflorescence a rusty tomentellous panicle 41. S. odoratissima 

19. Inflorescence a spike or fascicle. 

20. Leaves 8-21 cm. Reticulation faintly prominent. Inflorescence a fascicle. Bracts persistent. 

Fruits 10-12 mm long. Seeds straight. Stone different from the following . . 5. S. atjehensis 

20. Leaves 1 5-45 cm. Reticulation much prominent. Inflorescence a (basally branched) spike (a 

cone in bud). Bracts caducous. Fruits 8-10 mm long. Seeds not straight. Stone with a transverse 

constriction at one side 48. S. rubiginosa 

17. Twigs hairy. 
21. Leaves distichous. 
22. Fruits ampulliform. Inflorescence a fascicle. Bracts persistent. Stone ampulliform. 

23. S. fasciculata 

22. Fruits not ampulliform. Inflorescence not a fascicle. Bracts caducous. Stone not ampulliform. 

33. S. laeteviridis 
21. Leaves spirally arranged. 

23. Upper side of leaves hairy 3. S. adenophylla 

23. Upper side of leaves glabrous. 

24. Bracts persistent. 

25. Fruits 10-12 mm long and 5-6 mm broad 5. S. atjehensis 

25. Fruits to c. 10 mm long. 
26. Stone ellipsoid. Leaf margin (and petiole) beset with closely spaced glands. 

3. S. adenophylla 
26. Stone ampulliform. Fruits 5-7 mm. 
27. Nerves more than 10 pairs. Inflorescence a (basally branched) spike. 

16-1. S. cochinchinensis ssp. cochinchinensis 

27. Nerves less than 10 pairs. Inflorescence a fascicle 23. S. fasciculata 

24. Bracts caducous. 
28. Inflorescence not a spike. 

29. Inflorescence a panicle. Seeds not straight 41. S. odoratissima 

29. Inflorescence a (basally branched) raceme. 

30. Angle of leaf base c. 90° 50 S. sumatrana 

30. Angle of leaf base 20-60° 47. S. robinsonii 

28. Inflorescence a (basally branched) spike or a cone. 

31. Fruits to c . 10 mm long; mesocarp fleshy (shrivelled when dry) or thin, coriaceous. 

32. Leaves longer than 15 cm. Fruits 1-celled. Angle of leaf base 20-40°. Stone with low ridges 

and a depression or transverse groove near the base. Seeds not straight . . 48. S. rubiginosa 



1977] Symplocaceae (Nooteboom) 233 

32. Leaves 6-14 cm. Fruits 3-celled. Angle of leaf base c. 90°. Stone with low not interrupted 
ridges or grooves or brain-like grooved. Seeds straight 50. S. sumatrana 

31. Fruits 22-40 mm long; mesocarp woody or corky. 

33. Nerves 10-13 pairs 15b. S. cerasifolia var. grandifolia 

33. Nerves 6-9 pairs 15. S. cerasifolia 

Malay Peninsula 

1. Midrib prominent on the upper surface. 
2. Twigs hairy. 
3. Leaves evenly distributed. Fruits ellipsoid 4. S. anomala 

3. Leaves crowded towards the end of the twigs. Fruits ovoid 37. S. wikstroemifolia 

2. Twigs glabrous. 

4. Leaves crowded towards the end of the twigs. Fruits ovoid 37. S. wikstroemifolia 

4. Leaves evenly distributed. Fruits ellipsoid 35. S. lucida 

1. Midrib impressed on the upper surface. 
5. Twigs hairy. 
6. Leaves distichous. 
7. Inflorescence a raceme or panicle. Fruits ovoid to ellipsoid, 7-12 mm. Bracts caducous. 

33. S. laeteviridis 

7. Inflorescence a fascicle. Fruits ampulliform. Bracts persistent 23. S. fasciculata 

6. Leaves spirally arranged. 

8. Upper side of leaves hairy. 

9. Angle of leaf base more than 90°. Bracts caducous. Hairs on twigs more than 2 mm long. 

13. S. calycodactylos 
9. Angle of leaf base less than 90°. 
10. Fruits 8-10 mm long. Leaf margin (and petiole) beset with closely spaced glands. 

3. S. adenophylla 
10. Fruits more than 10 mm long. Leaf margin (and petiole) often glandular but glands not closely 

spaced 20. S. crassipes 

8. Upper side of leaves glabrous. 
11. Leaves crowded towards the end of the twigs. Inflorescence a spike. Fruits ovoid, 10-12 mm. 

37. S. wikstroemifolia 
11. Leaves evenly distributed. 
12. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular 
glands: 3. S. adenophylla.) 

13. Fruits cylindrical. Inflorescence a fascicle 28. S. glomerata 

13. Fruits ampulliform or ovoid. Inflorescence a spike or a panicle. 
14. Fruits ampulliform. Inflorescence a (basically branched) spike. Seeds not straight. 

16-1. S. cochinchinensis ssp. cochinchinensis 

14. Fruits ± ovoid. Inflorescence a panicle. Seeds curved 41. S. odoratissima 

12. Underside of leaves hairy (or pulverulent, nearly glabrous, in S. adenophylla). 
15. Bracts caducous. 
16. Stone smooth. Fruit 5-8 mm, ellipsoid .... 42-lb. S. ophirensis var. densireticulata 
16. Stone with ridges or grooves. 

17. Fruits 3-celled with 8 high ridges, 22^40 mm 15. S. cerasifolia 

17. Fruits 1 -celled. 

18. Inflorescence a panicle 41. S. odoratissima 

18. Inflorescence a (basally branched) spike (a cone in bud) 48. S. rubiginosa 

15. Bracts persistent. 

19. Fruits more than 13-18 mm long 20. S. crassipes 

19. Fruits to c. 10 mm long. 

20. Stone ellipsoid. Leaf margin (and petiole) beset with closely spaced glands 3. S. adenophylla 
20. Stone ampulliform. 
21. Nerves more than 10 pairs. Inflorescence a (tastily branched) spike. Seeds not straight. 

16 1. S. cochinchinensis up. cochinchinensis 

21. Nerves 6-8 pairs. Inflorescence a fascicle 23. S. fasciculata 

5. Twigs glabrous. 
22. Underside of leaves hairy. 
23. Leaves crowded towards the end of the twi 8 10 pairs. Stone smooth. Inflorescence a 

spike 37. S. wikstroemifolia 

23. Leaves evenly distributed or nerves more than 10 pans Stone with ridges or gTOOVM. 

24. Leaves distichous. Inflorescence a raceme or panicle 13, S. Iaete\iriilis 

24. Leaves spirally arr.m. 
25. Nerves less than 10 i tf persistent I mil. M IS mm long. Stone with low not inter- 

rupted rid oi brain-like gr light . 20. s. crassipes 

25. Nerves 12 17 pa I mils k 10 mm long stone with low ridgci and a depict* 

sion or transverse groove near the base Seeds not straight 48. S. ruhi^inosa 



234 Flora Malesiana [ser. I, vol. 8 2 

22. Underside of leaves glabrous. 

26. Leaves distichous 33. S. laeteviridis 

26. Leaves spirally arranged. 

27. Inflorescence terminal 46. S. pyriflora 

27. Inflorescence axillary. 
28. Bracts caducous. 

29. Inflorescence a spike. Fruits 25-40 mm 7. S. barringtoniifolia 

29. Inflorescence not a spike. Fruits shorter than 25 mm. 
30. Inflorescence a panicle. 

31. Petiole 7-10 mm 39. S. nivea 

46. S. pyriflora 

31. Petiole 10-50 mm 41. S. odoratissima 

30. Inflorescence a (basally branched) raceme (rarely a cone in bud). 

32. Fruits 3-celled 14. S. celastrifolia 

32. Fruits 1 -celled. 

33. Nerves 5-8 pairs. Terminal buds hairy 42. S. ophirensis 

51. S. sumuntia 

33. Nerves 9-14 pairs. Terminal buds glabrous 46. S. pyriflora 

28. Bracts persistent. 
34. Fruits ovoid to cylindrical. 

35. Acumen shorter than 5 mm. Apex of leaves rounded or acute 42. S. ophirensis 

35. Acumen longer than 5 mm. 
36. Nerves 10-16 pairs. Reticulation coarse. Inflorescence a fascicle. Fruits 7-10 mm long. 

28. S. glomerata 
36. Nerves 3-11 pairs. Reticulation fine. Inflorescence a (basally branched) spike. Fruits 13-18 

mm long 20. S. crassipes 

34. Fruits ampulliform. 

37. Petiole 2-9 mm. Inflorescence a raceme or panicle 42. S. ophirensis 

37. Petiole 10-15 mm. Inflorescence a spike 16-2. S. cochinchinensis ssp. laurina 



Java & The Lesser Sunda Islands 

1. Midrib prominent on the upper surface 35. S. lucida 

1. Midrib impressed in the upper surface. 
2. Twigs hairy. 

3. Leaves distichous 23. S. fasciculata 

3. Leaves spirally arranged. 
4. Underside of leaves glabrous. 
5. Fruits 8-25 mm 41. S. odoratissima 

5. Fruits 5-7 mm 16-1. S. cochinchinensis ssp. cochinchinensis 

4. Underside of leaves hairy. 

6. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

6. Upper side of leaves glabrous. 

7. Seeds not straight. Embryo at least U-shaped curved. 
8. Fruits ampulliform. Inflorescence a (basally branched) spike. Bracts persistent. Seeds twice 
curved 16-1. S. cochinchinensis ssp. cochinchinensis 

8. Fruits not ampulliform. Inflorescence a panicle. Bracts caducous. Seeds U-shaped 

41. S. odoratissima 
7. Seeds straight. Embryo at most slightly curved. 

9. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone not 
ampulliform 3. S. adenophylla 

9. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. Stone 

ampulliform 23. S. fasciculata 

2. Twigs glabrous. 

10. Fruits ampulliform 16. S. cochinchinensis 

10. Fruits spindle-shaped or otherwise not ampulliform. 
11. Inflorescence a (basally branched) spike or a cone. 
12. Angle of leaf base less than 60°. Fruits 20-40 mm long 19. S. costata 

12. Angle of leaf base more than 60°. Fruits to c. 10 mm long. 

16-4. S. cochinchinensis ssp. leptophylla 
11. Inflorescence not a spike (rarely a cone in bud). 

13. Terminal buds large (7-10 mm) 32. S. junghuhnii 

13. Terminal buds small. 

14. Inflorescence a panicle. Embryo curved 41. S. odoratissima 

14. Inflorescence a (basally branched) raceme. Embryo straight 11. S. brandisii 






1977] Symplocaceae (Nooteboom) 235 

Borneo 

1. Midrib flat or prominent on the upper surface 4. S. anomala 

1. Midrib impressed in the upper surface. 
2. Twigs glabrous. 
3. Underside of leaves hairy. 

4. Leaves distichous. Seeds straight 33. S. laeteviridis 

4. Leaves spirally arranged. 

5. Leaves 4-6 cm. Nerves 6-9 pairs. Petiole 3^* mm 10. S. brachybotrys 

5. Leaves 7^45 cm. Nerves 5-17 pairs. Petiole 10-50 mm. 

6. Leaves 15-45 cm. Nerves 12-17 pairs. Inflorescence a spike 48. S. rubiginosa 

6. Leaves 7-40 cm. Nerves 5-16 pairs. Inflorescence a panicle 41. S. odoratissima 

3. Underside of leaves glabrous. 

7. Leaves distichous 33. S. laeteviridis 

7. Leaves spirally arranged. 
8. Bracts persistent. 

9. Leaves shorter than 5 cm 12. S. buxifolia 

9. Leaves longer than 5 cm. 

10. Inflorescence not a spike 42. S. ophirensis 

10. Inflorescence a (basally branched) spike or a cone. 

11. Petiole 3-10 mm. Leaves 6-22 mm. Nerves 6-13 pairs 42. S. ophirensis 

11. Petiole 10-25 mm. 
12. Petiole 10-15 mm. Leaves 4 1 2 -21 cm. Nerves 6-9 pairs. 

16-2. S. cochinchinensis ssp. laurina 

12. Petiole 15-25 mm. Leaves 17-30 cm. Nerves 11-13 pairs 29. S. goodeniacea 

8. Bracts caducous. 
13. Inflorescence a (basally branched) spike or a cone in bud. 

14. Petiole 3-4 mm 10. S. brachybotrys 

14. Petiole more than 5 mm. (When dubious under "15", look under 53. 5. tricoccata.) 
15. Leaves longer than 15 cm. Nerves more than 10 pairs. Twigs thick. Terminal buds hairy. 

Petiole more than 20 mm. Fruits 25-40 mm 7. S. barringtoniifolia 

15. Leaves shorter than c. 10 cm. Nerves less than 10 pairs. Twigs not thick. Terminal buds 

glabrous. Petiole 5-10 mm 25. S. gambliana 

13. Inflorescence not a spike (rarely a cone in bud). 
16. Inflorescence a short spike, fascicle or panicle. 
17. Fruits 1-celled. Inflorescence a panicle. Stone with ridges or grooves. Seed 1, not straight. 

41. S. odoratissima 

17. Fruits 3-celled. Inflorescence a fascicle. Stone smooth. Seeds more than 1, straight. 

53. S. tricoccata 
16. Inflorescence a (basally branched) raceme. 

18. Leaves l 1 2 -5 cm. Nerves 6-9 pairs. Fruits ellipsoid, 10-15 mm long . . 12. S. buxifolia 
18. Leaves longer than 5 cm. 

19. Apex of leaf rounded to faintly acuminate. 
20. Leaves 5-8 cm. Nerves 6-9 pairs. Petiole 3-4 mm. Fruit unknown 9. S. borneensis 

20. Leaves 6-18 cm. Nerves 6-13 pairs. Petiole 3-10 mm. Fruit ovoid to ellipsoid, 5-12 mm. 

42. S. ophirensis 
19. Apex of leaf rather abruptly acuminate. 

21. Fruit globose, 3-celled, the sterile cells filled with air 14. S. celastrifolia 

21. Fruit ovoid to ellipsoid, 1-celled 42. S. ophirensis 

2. Twigs hairy. 
22. Leaves distichous. 

23. Underside of leaves glabrous 33. S. laeteviridis 

23. Underside of leaves fiairy. 
24. Leaf base 90-130'. Flowers solitary. Petiole 1-3 mm. Leaves ovate 2 l 2 -7 cm. Nerves 3-6 pairs. 

Fruit narrowly flask-shaped, c. 13 mm long 31. S. johniana 

24 These characters not combined. Fruits ellipsoid to ovoid. 
25. Petiole 3-4 mm. Leaves 4-12 cm. Nerves 7-11 pairs. Base of leaves 40-90 . Inflorescence more 
flowered. Bracts caducous. 

26. Leaves 4 9 cm, acumen 9 16 mm, base 40 90 17. S. colomrx.nensis 

2(> 12 cm, acumen 7 12 mm, base c. 90 33. S. laeteviridis 

25. Petiole ' , 3 mm, if up to 4 mm, inflorescence 1 -flowered. Nci rs. 

27. Acumen I 3 mm. Inflorescence 1-llowcred. Leaves 2 3' ,cm Hracts persistent. 

52. S. trichomarginalis 
27. These characters not combined 
28. Angle of leaf h. ; I lowers solitary or in a few-flowcred raceme. Hracts persistent, 

le I 2 mm. Leaves 2' . s ' 2 cm. Nerves 5 9 pairs. Acumen c. 5 mm. 

5K. S /i/vphuidt s 
28. These characters not combined 33. S. luttcwridis 

22. Leaves spirally arranged. 



236 Flora Malesiana [ser. I, vol. 8 2 

29. Leaves crowded towards the end of the twigs. Twigs thick, with large leaf-scars. 

44. S. polyandra 
29. Leaves evenly distributed. 
30. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular 
glands: 3. 5. adenophylla.) 
31. Inflorescence a spike. 

32. Leaves 4-6 cm. Petiole 3-4 mm 10. S. brachybotrys 

32. Leaves 6-25 cm. Petiole more than 5 mm. Nerves more than 10 pairs. Fruits ampulliform. Stone 

ampulliform 16-1. S. cochinchinensis ssp. cochinchinensis 

31. Inflorescence a panicle 41. S. odoratissima 

30. Underside of leaves hairy. 

33. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

33. Upper side of leaves glabrous. 
34. Seeds not straight. 

35. Petiole 3-4 mm 10. S. brachybotrys 

35. Petiole more than 5 mm. 
36. Fruits ampulliform. Bracts persistent . . . 16-1. S. cochinchinensis ssp. cochinchinensis 
36. Fruits not ampulliform. 

37. Inflorescence a panicle 41. S. odoratissima 

37. Inflorescence a (basally branched) spike (a cone in bud) 48. S. rubiginosa 

34. Seeds straight. 
38. Fruits to c. 10 mm long. 

39. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone 
ellipsoid 3. S. adenophylla 

39. Leaf index 2-3. Leaf margin (and petiole) often glandular but glands not closely spaced. 
Stone ampulliform 23. S. fasciculata 

38. Fruits more than 10 mm long. 

40. Angle of leaf base more than 60". 

41. Angle of leaf base less than 90'. Twigs and underside of leaves (appressedly) pubescent, 
puberulous or pilose. Fruits not ampulliform, 2-3-celled. Stone with ridges or grooves. 

20. S. crassipes 

41 . Angle of leaf base more than 90' . Twigs and underside of leaves not appressedly pubescent 
or puberulous. Fruits ampulliform, 1 -celled. Stone smooth 31. S. johniana 

40. Angle of leaf base less than 60'. 

42. Petiole 1-10 mm. Fruits 13-18 mm long 20. S. crassipes 

42. Petiole 1 5-25 mm. Fruits 22-40 mm long 15. S. cerasifolia 

Philippines 

1 . Leaves verticillate 55. S. verticillifolia 

1 . Leaves not verticillate. 
2. Midrib prominent on the upper surface. 
3. Twigs glabrous. Petiole more than 5 mm 35. S. lucida 

3. Twigs hairy. Petiole 1-5 mm 34. S. lancifolia 

2. Midrib impressed in the upper surface. 

4. Twigs hairy. 

5. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands: 

3. S. adenophylla.) 

6. Fruits spindle-shaped or otherwise not ampulliform. 

7. Leaves evenly distributed, 4 1 2-7*2 cm. Acumen 12-20 mm. Twigs not thick, cylindrical. Nerves 

5-6 pairs. Petiole 7-8 mm. Fruits 1-celled. Seed 1. Apex of leaves acuminate . . 24. S. filipes 

7. Leaves crowded towards the end of the twigs, longer than 10 cm. Acumen shorter than 5 mm. 

Twigs thick, tapering towards apex. Nerves more than 10 pairs. Petiole more than 20 mm. Fruits 

3-celled. Seeds more than 1. Apex of leaves rounded or acute 44. S. polyandra 

6. Fruits ampulliform 16-1. S. cochinchinensis ssp. cochinchinensis 

5. Underside of leaves hairy. 

8. Leaves distichous 23. S. fasciculata 

8. Leaves spirally arranged. 

9. Leaf margin and petiole beset with closely spaced glands 3. S. adenophylla 

9. Leaf margin and petiole not so. 
10. Fruits ampulliform. 
11. Nerves more than 10 pairs. Inflorescence a (basally branched) spike. 

16-1. S. cochinchinensis ssp. cochinchinensis 

11. Nerves less than 10 pairs. Inflorescence a fascicle 23. S. fasciculata 

10. Fruits spindle-shaped or otherwise not ampulliform. 

12. Petiole 1-5 mm 34. S. lancifolia 

12. Petiole more than 5 mm. 

13. Bracts and bracteoles caducous. Seeds not straight. Petiole 10-50 mm. 41. S. odoratissima 



1977] Symplocaceae (Nooteboom) 237 

13. Bracts and or bracteoles persistent. Seeds straight. Petiole 7-8 mm .... 24. S. filipes 

Petiole 5-7 mm 56. S. vidalii 

4. Twigs glabrous. 
14. Underside of leaves hairy 24. S. filipes 

14. Underside of leaves glabrous. 

15. Inflorescence a (basally branched) spike or a cone. 
16. Petiole 1-3 mm. Fruit ellipsoid to globose, 3-5 mm. Inflorescence a spike . 34. S. lancifolia 
16. Petiole longer than 3 mm. 

17. Petiole 15-25 mm. Inflorescence a spike to l 1 2 mm long 54. S. trisepala 

17. Petiole shorter than 15 mm or inflorescence longer than l 1 2 cm. 

18. Inflorescence a raceme or panicle of racemes. Embryo curved 42. S. ophirensis 

18. Inflorescence a spike. Embryo straight or curved. 

19. Inflorescence a short spike to 1\' 2 cm. Embryo straight 40. S. obovatifolia 

19. Inflorescence a spike, longer than V/ 2 cm. Embryo curved. 

16-1. S. cochinchinensis ssp. cochinchinensis 
15. Inflorescence not a spike (rarely a cone in bud). 
20. Fruits 2-5-celled. 

21. Stone with ridges or grooves 27. S. glabriramifera 

42. S. ophirensis 
21. Stone smooth. 
22. Acumen shorter than 5 mm. Inflorescence a fascicle to l l ' 2 cm. Bracts persistent. Fruits 

more than 10 mm long. Seeds straight 40. S. obovatifolia 

22. Acumen longer than 5 mm 14. S. celastrifolia 

20. Petiole 10-15 mm. Leaves 7-20 cm. Inflorescence a panicle. Fruit 8-25 mm, ovoid 

41. S. odoratissima 
20. These characters not combined. 

23. Leaves 2-5 3 ., cm. Fruits ovoid, 5-7 mm 57. S. whitfordii 

23. These characters not combined. 

24. Embryo straight. Leaves 4, l \ 2 -V\ 2 cm. Petiole 7-8 mm 24. S. filipes 

24. Embryo curved. Leaves 5-22 cm. Petiole 1-10 mm 42. S. ophirensis 

Celebes & The Moluccas 

1. Midrib prominent on the upper surface 35. S. lucida 

1. Midrib impressed in the upper surface. 
2. Twigs hairy. 
3. Leaves distichous. 

4. Underside of leaves glabrous 33. S. laeteviridis 

4. Underside of leaves hairy. 

5. Fruits ampulliform. Inflorescence a fascicle. Bracts persistent 23. S. fasciculata 

5. Fruits not ampulliform. Inflorescence not a fascicle. Bracts caducous . . . . 33. S. laeteviridis 
3. Leaves spirally arranged. 

6. Leaves crowded towards the end of the twigs 44. S. polyandra 

6. Leaves evenly distributed. 
7. Underside of leaves glabrous. (If leaf margin and petiole beset with closely spaced vesicular glands : 
3. S. adenophylla.) 
8. Seed and embryo twice curved 16-1. S. cochinchinensis ssp. cochinchinensis 

8. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 
7. Underside of leaves hairy. 

9. Upper side of leaves hairy (pulverulent) 3. S. adenophylla 

9. Upper side of leaves glabrous. 

10. Bracts caducous. 
11. Stone smooth. Inflorescence a (basally branched) raceme. 

42 lb. S. ophirensis fat (Knsireticulata 

11. Stone with ridges or grooves. Inflorescence a panicle of 5-30 cm . . 41. S. odoratissima 
10. Bracts persistent. 

12. Seeds not straight 16 4. S. cochinchinensis ssp. leptophylla 

12. Seeds straight. 

13. Leaf index more than 3. Leaf margin (and petiole) beset with closely spaced glands. Stone 

ellipsoid I s - adenophylla 

13. Leaf index 2 3. Leaf margin (and petiole) often glandular but glands MX closely spaced. 
Stone ampulliform .... 23. S. fasciculata 

2. Twigs glabr. 
14. Underside of leaves hairy. 

15. Leaves distichous. Petiole I 5 mm. Bracts cadii .lit 33. S. laeteviridis 
15. Leaves spirally arranged. 16. 4. S cochinchinensis s\p leptophylla 

14. Underside of leaves glabrous. 



238 Flora Malesiana [ser. I, vol. 8 2 

16. Fruits 2-3-celled. 
17. Leaves longer than 15 cm. Fruits more than 10 mm long. Leaf margin entire. Nerves 9-14 pairs. 
Stone with ridges or grooves. Seeds straight 36. S. maliliensis 

17. Leaves shorter than c. 15 cm. Fruits to c. 10 mm long. Seed and embryo U-shaped. 

14. S. celastrifolia 
16. Fruits 1 -celled. 

18. Leaves distichous 33. S. laeteviridis 

18. Leaves spirally arranged. 

19. Inflorescence a panicle of 5-30 cm 41. S. odoratissima 

19. Inflorescence not a panicle or shorter than 5 cm. 

20. Fruit stone with high, interrupted ridges which often protrude from the base. 

42. S. ophirensis 

20. Stone different 16. S. cochinchinensis 



New Guinea 
(incl. New Ireland & New Britain) 

1. Leaves (pseudo-)verticillate 30. S. herzogii 

1. Leaves not verticillate. 
2. Twigs glabrous. 

3. Underside of leaves hairy 16-4. S. cochinchinensis ssp. leptophylla 

3. Underside of leaves glabrous. 

4. Nerves 13-20 pairs. Leaves 21-62 cm 26. S. gigantifolia 

4. Nerves less than 15 pairs. Leaves usually much smaller. 
5. Twigs thick. 
6. Acumen shorter than 5 mm. Fruits c. 13 mm long. Nerves 8-12 pairs ... 45. S. pulvinata 

6. Acumen longer than 5 mm. Fruits to c. 10 mm long. 

16-4. S. cochinchinensis ssp. leptophylla 
5. Twigs not thick. 

7. Bracts persistent. 

8. Inflorescence a panicle to 5 cm. Petiole 13-15 mm. Nerves 5-9 pairs . 18. S. composiracemosa 

8. Plant different 16. S. cochinchinensis 

7. Bracts caducous. 

9. Fruits c. 15 mm long 21. S. cylindracea 

9. Fruits to c. 10 mm long. 

10. Fruits 3-celled (often 1 or 2 aborted) 14. S. celastrifolia 

10. Fruits 1-celled 16-3. S. cochinchinensis ssp. thwaitesii 

2. Twigs hairy. 
11. Underside of leaves glabrous. 
12. Inflorescence only 1-flowered. 

13. Leaves shorter than 5 cm 16-4. S. cochinchinensis ssp. leptophylla 

13. Leaves longer than 5 cm. 
14. Leaf index less than 2. Acumen shorter than 5 mm. Angle of leaf base less than 90°. Nerves less 
than 5 pairs. Reticulation coarse. Fruits to c. 10 mm long. Seeds not straight. Apex of leaves 

rounded or acute 16-4. S. cochinchinensis ssp. leptophylla 

14. Leaf index more than 2. Acumen longer than 5 mm. Angle of leaf base more than 90°. Nerves 
more than 5 pairs. Reticulation fine. Fruits 17-22 mm long. Seeds straight. Apex of leaves 

acuminate 38. S. multibracteata 

12. Inflorescence more-flowered. 

15. Petiole 0-5 mm 16-4. S. cochinchinensis ssp. leptophylla 

15. Petiole more than 5 mm. 

16. Seed and embryo (twice) curved 16-1. S. cochinchinensis ssp. cochinchinensis 

16. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 
1 1 . Underside of leaves hairy. 

17. Upper side of leaves hairy 43. S. paucistaminea 

17. Upper side of leaves glabrous. 
18. Inflorescence only 1-flowered. 
19. Leaf index more than 3. Angle of leaf base less than 90°. Reticulation coarse . 49. S. salicioides 

19. Leaf index 2-3. Angle of leaf base more than 90°. Reticulation fine . . 38. S. multibracteata 
18. Inflorescence more-flowered. 

20. Bracts caducous. Seeds straight 15. S. cerasifolia 

20. Bracts persistent. Seeds not straight. 

21. Seed and embryo (twice) curved 16-1. S. cochinchinensis ssp. cochinchinensis 

21. Seed and embryo uncinately curved towards the base. 

16-4. S. cochinchinensis ssp. leptophylla 



1977] 



Symplocaceae (Nooteboom) 



239 



3. Symplocos adenophylla Wall. (Cat. 1831, 
n. 4427A, nomen) ex G. Don, Gen. Syst. 4 (1837) 3 ; 
DC. Prod. 8 (1844) 257; Miq. Fl. Ind. Bat. 1, 2 
(1859) 466; Clarke, Fl. Br. Ind. 3 (1882) 575; 
Brand, Pfl. R. Heft 6 (1901) 48, incl. var. virgata 
Wall. (Cat. 1831, n. 4427B, nomen) ex Brand; 
K. & G. J. As. Soc. Beng. 74, ii (1906) 240; Brand, 
Bull. Herb. Boiss. II, 6 (1906) 747, incl. var. atrata 
Brand, I.e. 748; Merr. Philip. J. Sc. 2 (1907) Bot. 
298; Brand, Philip. J. Sc. 3 (1908) Bot. 7, incl. var. 
merrittii Brand; Ridl. Fl. Mai. Pen. 2 (1923) 303, 
t. 101, incl. var. montana Ridl.; Noot. Leid. Bot. 
Ser. 1 (1975) 121. — S. bancana Miq. Fl. Ind. 
Bat. Suppl. 1 (1861) 476. — S. iteophylla Miq. I.e., 
incl. var. rostrata Miq. et var. elliptica Miq. ; Merr. 
En. Born. (1921) 486. — Eugeniodes adenophyllum 
O. K. Rev. Gen. PI. 2 (1891) 410. — S. beccarii 
Brand, Pfl. R. Heft 6 (1901) 49. — S. const ricta 
Brand, I.e. 41; Merr. En. Born. (1921) 486. — 
S.fulvosa King & Gamble, J. As. Soc. Beng. 74, ii 
(1906) 233; Ridl. Fl. Mai. Pen. 2 (1923) 300. — 
S. palawanensis Brand, Philip. J. Sc. 3 (1908) Bot. 
10; Merr. En. Philip. 3 (1923) 301. —S.pruniflora 
Ridl. J. Fed. Mai. St. Mus. 4 (1909) 46; Fl. Mai. 
Pen. 2 (1923) 304. — S. brandii Elmer, Leafl. 
Philip. Bot. 4(1912) 1477. — S. pahangensis Brand 
in Fedde, Rep. 14 (1916) 326. — Fig. 7. 

Shrub or tree to 20 m, 50 cm 0. Young twigs 
pulverulent-puberulous or rarely tomentellous, 
glabrescent, often dark-brown to blackish. Inno- 
vations light redbrown. Leaves chartaceous to 
coriaceous, often dark brown when dry, pulverulent 
beneath or on both faces, soon glabrescent, elliptic, 
acuminate, with cuneate base and recurved to 
revolute margin with many pellucid glands, 4 1 /}- 
16 by l 1 4 -4 3 / 4 cm; nerves 4—12 pairs, meeting in a 
looped intramarginal vein; petiole 6-12 mm. 
Flowers in a spike, raceme or panicle to 6 cm; in- 
dument of axis as twigs. Bracts and bracteoles with 
same indument persistent in fruit, l /j-l mm. Pedicel 
mostly only under older flowers, to 3 mm. Calyx 
nearly entirely divided into '/j-l mm long lobes. 
Corolla 2-5 mm. Stamens (20-)25-50. Disk gla- 
brous or rarely hairy. Ovary with same indument as 
that of twigs, 1-2 mm high; style glabrous or with 
some hairs towards the base, 2-4 mm. Fruit ellip- 
soid to cylindrical, sometimes with c. 6 ridges when 
dry, blue or black-purple, soon glabrescent, 
crowned by the incurved calyx lobes, with only one 
developed cell, 8-10(-ll) by 3-5(-6) mm. Seed 1, 
with straight embryo. 

Distr. Continental Asia (China incl. Hainan, 
Indo-China, Thailand), throughout Malesia, except 
Java (but found in Bawean I.), the Lesser Sunda Is. 
and New Guinea. A variety in Indo-China. 

Ecol. Usually in montane rain-forest, in moun- 
tain heaths, on ridge-crests and ridges, and mossy 
forest, also in Baeckea-Lep/ospcrmum heath forest, 
often on granite, but also on ultra basic ( I rusmadi), 
from sea-level to 3000 m, but at low altitude 
largely Of] podsottssd sand (Manka; Baku N. P.) 
and in heath hues! on humid podsol. / /. Sept. 
(lebr. Oct.),/r. May (Jan. Dec). 

As is the case with more species, dwarfed 

•r hardly I m high may already COfM Into 
flower. 

Uses. The timber can be used for light construc- 
tions n>i s< h. Mai, i or. i<ec. IV, 1954, 593). 

Vcrn. & Uses. MCndong, mtnugan, Malaya, 



kaju lattan, k. porugis, Sumatra, Batak, kayu kain, 
W. Borneo, G. Klamm; the latter name alluding 
to the use for tanning cloth in dyeing. 

The Besisi (Mai. Pen.) believe that the leaves of 
certain plants, e.g. S. adenophylla, if carried in the 
quiver with their darts, act as charms bringing them 
success in hunting (Burk. Diet. 1935). 

4. Symplocos anomala Brand, Bot. Jahrb. 29 
(1900) 529; Pfl. R. Heft 6 (1901) 67; Noot. Leid. 
Bot. Ser. 1 (1975) 126, pi. la-f, with full synonymy. 
— 5. concolor Brand, Pfl. R. Heft 6 (1901) 65; 
K. & G. J. As. Soc. Beng. 74, ii (1906) 242; Ridl. 
Fl. Mai. Pen. 2 (1923) 304. — Fig. 7. 

Shrub or tree to 21 m, 40 cm 0. Young twigs 
tomentellous to tomentose or appressedly pubes- 
cent, glabrescent. Leaves glabrous, brownish or 
olive to yellowish green glossy above, elliptic, 
acuminate with cuneate-attenuate base and more 
or less revolute finely glandular dentate to nearly 
entire margin, 2 1 / 2 -12 by l l /4-3 cm; midrib promi- 
nent above or flat, rarely flat and sunken; nerves 
5-1 1 pairs, meeting in a looped intramarginal vein; 
petiole 2-7 mm. Raceme to 2 cm long, axis tomen- 
tose to appressedly pubescent. Bracts 1-2 mm, 
bracteoles 1 U-V[ 2 mm, both persistent, with same 
indument as axis. Pedicels 2-5 mm. Calyx lobes 
rounded, ciliolate l l 2 -2 mm. Corolla 4-6 mm. 
Stamens 50 to more than 100. Disk tomentose or 
shortly soft hairy. Ovary tomentose to (finely) 
appressedly pubescent, c. '^-IVjmm high; style 
glabrous or hairy towards the base, 4-7 mm. Fruit 
3-celled, ellipsoid, violet, almost black, c. 10 by 
6 mm in Malaya, 10-13 by 6-8 mm in Borneo. 
Seed 1 in each cell, straight with straight embryo. 

Distr. Continental Asia (Burma, Thailand, 
Indo-China, China incl. Hainan, Japan, Ryu 
Kyu Is., Formosa) and Malesia: Malaya (incl. 
Penang), N. Sumatra (incl. Banka), and Borneo. 

Ecol. Mixed evergreen montane forest, also on 
ridges and along streamsides, 700-2200 m (in 
continental Asia to 3000 m), but also found on 
podsolized sands at very low altitude, 20-50 m, in 
Banka. Fl. June-Oct.,/r. Jan.-Dec. 

Vern. Rinak, Banka. 

5. Symplocos atjehensis Noot. Leid. Bot. Ser. 1 
(1975) 128. — Fig. 7, 8. 

Treelet to r. 8 m, 10 cm 0. Twigs glabrous or 
tomentose. Leaves glabrous or sparsely appressedly 
hairy, especially on midrib and nerves, elliptic, 
acuminate, with acute to rounded base and dentate 
margin, 8-21 by 3'/ 2 -6cm; nerves 8-12 pairs, 
meeting in a looped intramarginal vein; petiole 
12-17 mm. I lowers in fascicles with persistent red- 
bum n tomentoac to pubeecent e. 2 mm long bracts 
and bracteoles. Calyx 2 mm, the (ciliatc) lobes 
I |'/j mm. Corolla c. 5 mm. Stamens c. 50. Disk 

glabrous. Ovary glabroui without, <■. 1 mm high; 
itylc glabrous, A 6mm. Fruit ellipsoid, 10 12 by 

J 6mm, 1-ccllcd. but only one cell developing; 
stone shallow ly lengthwise nhhed. Seed I, straight 
With straight embryo. 

Distr. Mulesui: N. Siimalia ((iajo I anils). 

1 col. Mixed eve r g r e en mountain forest, 1700 

IHSOm. II. Aug. Sept../r. July. 

<>. Symplocos burisunicu N001. leid. Hot. Ser. 1 
(1975) HO. lig. 7. 



240 



Flora Malesiana 



[ser. I, vol. 8 2 




**«-TSTjrnt»casaasflfia5Mr'--— 






1977] 



Symplocaceae (Nooteboom) 



241 







Fig. 9. Synpbcos bataketuis Noon .1 Habit, nat. size. S. barrtngtonilfolta Brand b. Fruit, c ditto in 

( S, both I 1 , (fl Kmhinson & Ki.oss 125, b 1 Kll'l Kl 10736). 



242 



Flora Malesiana 



[ser. I, vol. 8 2 



Small tree, 6-10 m, 25 cm 0. Twigs glabrous. 
Leaves glabrous, elliptic, with acute to nearly 
rounded base, denticulate margin and acuminate to 
rounded apex, 12-20 by 5-12 cm; nerves 8-12 
pairs, whether meeting in an intramarginal vein or 
not; petiole 10-30 mm. Spike branched, to 5 cm 
with minutely appressedly hairy axis. Bracts and 
bracteoles persistent, ± ovate, with same indument, 
2-3 mm long. Calyx divided into semi-orbicular, 
glabrous but ciliolate 1 '/ 2 -2 mm long lobes. Corolla 
c. 6 mm. Stamens c. 50. Disk glabrous. Ovary c. 
1 mm high, glabrous; style glabrous, 4-5 mm, or 
reduced. Fruit ovoid to globose, c. 10 by 9 mm with 
globose to ampulliform stone of c. 8 by 7 mm (the 
neck c. 2 mm long and the belly irregularly length- 
wise grooved, c . 6 mm high). Seed 1 , U-shaped 
with U-shaped embryo. 

Distr. Malesia: Central W. Sumatra (Mts 
Kerintji and Merapi). 

Ecol. Montane rain-forest, on Mt Kerintji in 
Gleichenia woodland, 2000-2600 m. Fl. June-July. 

7. Symplocos barringtoniifolia Brand, Ann. Cons. 
Jard. Bot. Geneve 4 (1904) 283; Noot. Leid. Bot. 
Ser. 1 (1975) 131, pi. 4. — Doxomma rigidum 
Miers, Trans. Linn. Soc. II, Bot. 1 (1875) 104. — 
Barringtonia rigida Clarke, Fl. Br. Ind. 2 (1879) 
510. — S. rigida Clarke, Fl. Br. Ind. 3 (1882) 581, 
non G. Don, 1837; Brand, Pfl. R. Heft 6 (1901) 
52; K. & G. J. As. Soc. Beng. 74, ii (1906) 246; 
Ridl. Fl. Mai. Pen. 2 (1923) 306. — Eugeniodes 
rigidum O. K. Rev. Gen. PI. 2 (1891) 976. — Fig. 7, 
9b-c. 

Tree to 25 m, 40 cm 0. Twigs glabrous, often 
marked with prominent orbicular scars of fallen 
leaves; growth discontinuous, terminal buds pro- 
tected by leathery scales, leaving conspicuous scars. 
Leaves glabrous, elliptic to obovate with cuneate 
base and acuminate apex, 15-35 by 6-11 cm; 
nerves 10-14(— 16) pairs; petiole 2-5 cm. Spike 
resembling a cone in bud because of the large 
bracts, becoming 572(-8)cm; axis tomentose. 
Bracts and bracteoles tomentellous to appressedly 
pubescent, both soon caducous, broadly ovate, 
6-10 by 6 mm and narrowly ovate, 2 l / 2 -5 mm long 
respectively. Calyx tomentellous, 3-37 2 mm long, 
the 5 lobes originally c. 1 mm long but the calyx 
becoming 2-3-lobed by tearing. Corolla 4-6 mm. 
Stamens c. 60 to more than 100. Disk glabrous. 
Ovary glabrous, I-IV2 rnrn high; style c. 5 mm, 
with soft hairy conical base. Fruit ovoid or ellipsoid, 
royal blue, 272-4 by 1 72-2 cm, with chartaceous 
mesocarp; stone stellate in cross-section with 8 
very high ridges; cells 3, often only 1 fertile. Seed 
straight with straight embryo. 

Distr. Continental Asia (Indo-China), in 
Malesia: Malay Peninsula and Borneo (only once: 
W. Kutei). 

Ecol. Lowland rain-forest, river valleys in low 
undulating country, on hillsides on clay, on dry 
hillocks in Dryobalanops forest, but also on sand- 
stone or granite, mostly below 300 m, but also in 
Malaya more rarely in montane forest up to 
1500 m. Fl. July-Aug.,/r. Febr.-May (July). 

Vern. Medang, Malaya. 

8. Symplocos batakensis Noot. Leid. Bot. Ser. 1 
(1975) 132.— Fig. 7, 9a, lOa-d. 

Twigs glabrous. Leaves often coriaceous, glab- 



rous, elliptic (to obovate) with acute base, entire 
margin and acuminate apex, 6-10 by 2-472 cm; 
nerves 7-10 pairs, meeting in an intramarginal 
vein; petiole 5-8 mm. Raceme to 8 cm, axis glab- 
rous or sparsely minutely pilose. Bracts and brac- 
teoles with same indument, ovate, caducous, P/2 
and 1 mm respectively. Pedicel to 2(-5) mm. 
Calyx glabrous, 172 mm long, the semi-orbicular 
lobes 1-1 74 mm long. Corolla c. 6 mm. Stamens 
c. 100. Disk shortly pilose. Ovary glabrous, P/2 mm 
high ; style glabrous, c. 5 mm, sometimes reduced. 
Fruit nearly globose, c. 10 by 8 mm, or ellipsoid- 
ampulliform, c. 10 by 5 mm, 3-celled, often only 1 
cell fertile. Seed often only 1, straight with straight 
embryo. 

Distr. Malesia: Central W. Sumatra (Tapanuli 
and Westcoast Res.). 

Ecol. Montane rain-forest on low ridges, 1200- 
1700 m. Fr. Jan., Aug. 

Vern. Loala lola, sihondung, Tapanuli. 




Fig. 10. Symplocos batakensis Noot. a. Flowers and 
buds, b. fruit, c. endocarp, d. ditto in CS, all x 2. 
— S. cerasifolia Wall, ex DC. e. Fruit in CS, nat. 
size (a-d Robinson & Kloss 125, e SAN 45168). 



9. Symplocos borneensis Brand, Pfl. R. Heft 6 
(1901) 56; Merr. En. Born. (1921) 486; Noot. 
Leid. Bot. Ser. 1 (1975) 134. — Fig. 7. 

Twigs glabrous. Leaves narrowly elliptic, glab- 
rous, with acute base, entire margin and rounded 
to faintly acuminate apex (the acumen with broad 
rounded tip), 47 4 -8 by P/4-272cm; nerves 6-9 
pairs, faintly prominent beneath, meeting in an 
intramarginal vein ; reticulation hardly prominent ; 



1977] 



Symplocaceae (Nooteboom) 



243 



petiole 3-4 mm. Raceme lax, to 5 cm, axis minutely 
sparsely hairy to glabrous. Bracts and bracteoles 
caducous, glabrous, ciliolate, c. 1 and c. l /a mm 
long respectively. Pedicel to 2 mm. Calyx glabrous, 
divided into 3 4 -l mm long ciliolate lobes. Corolla 
c. 5 mm. Stamens 60-80. Disk shortly pilose. 
Ovary glabrous, l-l' 4 mm high; style glabrous 
except the conical shortly pilose base, c. 5 mm long. 
Fruit unknown. 

Distr. Malesia: Borneo (Sarawak and W. Bor- 
neo: Kenepai), 2 collections. 

Ecol. Lowland rain-forest. 

Note. A sterile collection from Central Celebes 
(Malili) possibly belongs to this species. 

10. Svmplocos brachybotrys Merr. J. Str. Br. R. 
As. Soc. n. 76 (1917) 110; En. Born. (1921) 486; 
Heine, Pfl. Samml. Clemens (1953) 87; Noot. Leid. 
Bot. Ser. 1 (1975) 134. — Fig. 7. 

Twigs (sparsely) appressedly pubescent in inno- 
vations, soon glabrescent. Leaves sparsely appres- 
sedly fine hairy when young, soon glabrescent, 
ovate to elliptic, with acute to rounded base, 
denticulate margin and acute to acuminate apex, 
4-6 by 2-3' 4 cm; nerves 6-9 pairs, meeting in an 
intramarginal vein; petiole 3-4 mm. Spike short, 
1-3-flowered, axis at most 7 mm, appressedly 
pubescent, or flowers solitary, sessile from the leaf 
axils and then several appressedly pubescent 2-4 
mm long bracts. Bracts and bracteoles caducous, 
in the spikes not seen. Calyx appressedly pubes- 
cent, divided into the c. 3 mm long lobes. Corolla 
6 mm. Stamens c. 100. Disk glabrous, inconspi- 
cuous. Ovary appressedly pubescent, 1 mm high; 
style glabrous, c. 4 mm. Fruit ovoid to ellipsoid, 
intense indigo-blue, c. 10 by 5 mm, stone shallowly 
lengthwise grooved. Seed 1, ovoid, slightly curved 
with S-shaped embryo. 

Distr. Malesia: Borneo (Sarawak and Sabah: 
Mt Kinabalu). 

Ecol. Mixed, evergreen mountain forest, 
1500-1800 m. 

11. Symplocos brandisii K. & V. Bijdr. 7 (1900) 
157; Brand, Pfl. R. Heft 6 (1901) 90; Koord. 
Atlas 2 (1914) t. 381 ; Back. & Bakh./. Fl. Java 2 
(1965) 206; Noot. Leid. Bot. Ser. 1 (1975) 135, 
pi. 5. — S. koordersiana Brand, Bull. Herb. Boiss. 
II, 6 (1906) 748. — S. pseudoclethra Hall./. Med. 
Rijksherb. 14(1912)41. 

Tree to 30 m, 40 cm 0. Twigs glabrous. Leaves 
glabrous, mostly narrowly elliptic, with attenuate 
base, (coarsely) crenate margin and hardly acumi- 
nate apex, (5 1 1 )7 13( 22) by 2' , 5( '(,> ,)cm; 
ner.es (7 )I0 ]f>( IK) pairs, meeting in a looped 
intramarginal vein; petiole 6-15 mm. Raceme to 
10 cm, but often shorter, axis (sparsely) pubescent. 
Bracts very soon caducous, appressedly pubescent, 
obovate or elliptic, I 4 mm ; bracteoles falling altei 
the bracts, sometimes rather long persistent, less 
hairy, (broadly) ovate to narrowly elliptic, I 2 1 , 
mm. Pedicels pubescent, at most 6 mm but often 
shorter. Calyx glabrous, or some appressed h.ius 
on the base of the tube, l*/j 2' ,. mm lone-, the 
lobes ft ' « mm shorter, sometime, ciliolate, 
Corolla ( . A( 5) mm. Stamens (A) to mOTC tli.m MX) 
Disk 5-glandular, glahrous. Ovar) fjabfOUS, 
I I ' .. mm hi^h; style glabrous or with lew bain, 
but the conical base soft-hairy, 4 5mm In/it 



ovoid to ellipsoid, slightly narrowed towards the 
apex, 10-16 by 5-7 mm; stone 1-celled, smooth or 
faintly ribbed. Seeds 1 (or 2), filling the whole 
stone, ovoid, with straight embryo. 

Distr. Malesia: Java and Lesser Sunda Is. 
(Lombok). 

Ecol. From sea-level to 1800 m. 



KEY TO THE VARIETIES 

1. Leaves 5V2-13cm. Bracts obovate, 1-3 mm. 

Stamens c. 60 a. var. brandisii 

I. Leaves 11-22 cm. Bracts elliptic, 3-4 mm. 

Stamens more than 100 . b. var. pseudoclethra 

a. var. brandisii. — S. brandisii K. & V. — S. koor- 
dersiana Brand. Cf. Noot. Leid. Bot. Ser. 1 (1975) 
136, pi. 5a-h. — Fig. 7. 

Leaves 5 l / 2 -13cm long. Nerves 7-16 pairs. 
Bracts obovate, 1-3 mm; bracteoles ovate to 
narrowly elliptic, l-2V2nim. Corolla c. 4 mm. 
Stamens c. 60. 

Distr. Malesia: West Java (Udjung Kulon 
Peninsula, Peutjang I. and Depok), East Java 
(Besuki: Pantjur Idjen), and Lesser Sunda Is. 
(Lombok: Mt Rindjani). 

Ecol. Lowland primary and secondary forest, in 
P. Peutjang on raised coral, in Java below 200 m, 
in Lombok in montane forest at '800-1800 m'. Fl. 
March-June, Nov.,//-. July. 

b. var. pseudoclethra (Hall. /.) Noot. Leid. Bot. 
Ser. 1 (1975) 136, pi. 5i-j. — S. pseudoclethra Hall. 
/ 

Leaves 11-22 cm. Nerves 11-18 pairs. Bracts 
elliptic, boat-shaped, 3-4 mm; bracteoles broadly 
ovate, c. 2 mm. Corolla l l / 2 mm. Stamens more 
than 100. 

Distr. Malesia: Lesser Sunda Is. (Lombok: Mt 
Rindjani). Only known from the type. 

Ecol. Montane high forest, 800-950 m. Fl. April. 

12. Symplocos buxifolia Stapf, Trans. Linn. Soc. 
Bot. 4 (1894) 206; Brand, Pfl. R. Heft 6 (1901) 64; 
Merr. En. Born. (1921) 487; Noot. Leid. Bot. 
Ser. 1 (1975) 136, pi. 6a-d. — Fig. 7. 

Shrub or treelet, 2 10 m; crown dense, globular, 
fastigiate. Twigs glabrous, dark, • zigzag. Leaves 
glabrous, closely placed, elliptic to nearly orbicular 
with more or less attenuate base, finely glandulai 
dentate or crenate margin and rounded to acute 01 
slightly acuminate apex, 15 50 by 7 25 mm; neives 
■J 6 pans, meeting in an intramargina] vein; petiole 
3-7 mm. Inflorescence an axillary few-flowered 

raceme Of often a 1 -(lowered shoot with several 

miniature sparsely pubescent to glabroiu bract' 

like leaves 01 3 by 1 to 10 by 5 mm ; a\is glabrous 01 

minutely appressedlj dairy. Bracts and bracteoles 
caducous; pedicel betwe en them to 2 mm. Calyx 
glabrous or finely appressedly hairy, - -1 S mm long, 

the lobes cihale, I 1 mm ( 'orolla s 8 mm. Stamens 

'ii to mora than 100. i>isk glabrous. Ovary glab 
ions oi rare!) find) appressedly hairy. 2 3mm 
high; style nabi mm. inm ellipsoid to 

ovoid, 10 I s by 6 Bmm; stone with low length' 
Seed i . .ii. hi'Iii with straight embryo. 
i > i s 1 1 \f,ii,M,i n. Borneo (Sabah: Ml Kina- 
balu). 



244 



Flora Malesiana 



[ser. I, vol. 8 2 



Ecol. Mixed, evergreen, subalpine low forest 
and scrub, common, 2400-4000 m.Fl. March-July, 
Oct., Dec.,/r. Febr.-Aug. 

Note. This species can hardly be distinguished 
from the mountain forms of S. cochinchinensis ssp. 
leptophylla in New Guinea, especially those with 
small orbicular leaves. 

13. Symplocos calycodactylos Brand, Pft R. 
Heft 6 (1901) 63; Noot. Leid. Bot. Ser. 1 (1975) 
137, pi. 6e. 

Shrub, 3 m. Twigs densely spreadingly long- 
hairy, hairs to 3 mm. Leaves long-hairy on both 
surfaces, ovate to elliptic with rounded to sub- 
cordate base, dentate, long-ciliate margin and 
acuminate apex, 6-14 by 2 l / 2 -5cm; nerves 7-8 
pairs, meeting in an intramarginal vein; petiole 
3-5 mm. Inflorescence a fascicle (or flowers soli- 
tary?) or raceme to 10 cm; axis long-hairy. 
Bracts and bracteoles soon caducous, to 7 mm long, 
narrowly elliptic clothed with long hairs. Pedicels 
from ?5 mm in fascicles to 13 mm in racemes. 
Calyx entirely divided into the narrow-elliptic to 
linear, 4-6 mm long pubescent lobes. Corolla c. 6 
mm. Stamens c. 100. Disk pilose. Ovary obscured 
by the 3 mm long hairs, l 1 ] 2 mm high; style glab- 
rous, c. 8 mm. Fruit ± cylindrical, densely long- 
hairy, crowned by the persistent calyx (only young 
fruits seen). 



Distr. Malesia: Malay Peninsula (Perak and 
Kedah), 2 collections. 

Ecol. Evergreen hill forest, 900-1000 m. Fl. 
Febr. 

Note. Closely allied to the Indian-Ceylonese 
S. pulchra Wight with which there are hardly any 
vegetative differences ; in flower easily distinguished 
by the extremely long calyx lobes. 

14. Symplocos celastrifolia Griff, ex Clarke, Fl. 
Br. Ind. 3 (1882) 575; Brand, Pfl. R. Heft 6 (1901) 
48; K. & G. J. As. Soc. Beng. 74, ii (1906) 239; 
Ridl. Fl. Mai. Pen. 2 (1923) 302; Merr. Un. Cal. 
Publ. Bot. 15 (1929) 248 ; Fletcher, Fl. Siam. En. 2 
(1938) 385; Noot. Leid. Bot. Ser. 1 (1975) 138. — 
Eugeniodes celastrifolius O. K. Rev. Gen. PI. 2 
(1891) 975. — S. nigricans Brand, Pfl. R. Heft 6 
(1901 ) 49. — 5. candicans Brand, I.e. — S. hutchin- 
sonii Brand, Philip. J. Sc. 4(1909) Bot. 109; Merr. 
En. Philip. 3 (1923) 299. — S. peninsularis Brand, 
Philip. J. Sc. 4 (1909) Bot. 110. — Fig. 4m, 7, 11. 
Shrub or small tree, rarely up to 30 m high and 
60 cm 0. Twigs glabrous. Leaves glabrous, or 
rarely sparsely fine-hairy on midrib and nerves 
beneath, often the upper surface dark coloured to 
nearly black when dry and the undersurface olive 
brown, ± elliptic, with cuneate-attenuate base, 
crenate margin and mostly abruptly acuminate 
apex, 5V 2 -15 by 2V 4 -6cm; nerves 6-9 pairs, 




Fig. 11. Symplocos celastrifolia Griff, ex Clarke, a. Habit, x 2 / 3 , b. bud, with bract and bracteoles, 

c. corolla and stamens, both x 3, d. anther, e. stigma, both x 9, f. LS of flower, x 5, g. CS of fruit, h. 

LS of fruit, both x 9 (a Main 1258, b-h Kostermans 1144, all from Morotai I.). 



1977] 



Symplocaceae (Nooteboom) 



245 



usually meeting in the intramarginal reticulation; 
petiole 3-15 mm. Raceme often basally branched, 
axis fine-hairy to appressedly pubescent, 3-12 cm. 
Bracts and bracteoles soon caducous, 2-3(-4 in 
Morotai) and c. P/a (° r 2-2 1 , in Morotai) mm 
long respectively. Pedicels with same indument as 
axis, 1-5 mm. Calyx glabrous, l 1 ■' 2 -2 1 2 (-3 in 
Morotai) mm; lobes ciliate, when young 1-1 ' 2 mm, 
becoming longer by tearing apart. Corolla 4-6 mm. 
Stamens 40-c. 60. Disk glabrous, with some hairs or 
pilose, especially after anthesis. Ovary glabrous, c. 
1 mm high; style glabrous, 4-5 mm. Fruit orbicular, 
pink, green, yellow or dark blue (sec. coll.), 4-10(- 
20) by 3-8(-15)mm; stone smooth, cells 3, but 
usually only 1 fertile, the sterile cells larger than the 
fertile ones, towards the base filled with air. Seed 
and embryo U-shaped. 

Distr. Peninsular Thailand and throughout 
Malesia, except in Java, the Lesser Sunda Is., the 
northern islands of the Philippines, the northern 
half of Celebes, and most of the Moluccas. The 
number of collections in Sumatra and East Malesia 
(E. of Makassar Straits) is small compared with 
those in Malaya and especially Borneo. 

Ecol. Usually in coastal, primary and secondary 
lowland forests especially in the transition zone 
between mangrove (Nypa) and freshwater swamps, 
mostly in deep marshy, sandy soils, but in a variety 
of other habitats: sandy beaches, sandbanks near 
the sea, kerangas, Casuarina peat swamp, in lalang 
fields on white sandy soils, open heath forest 
behind the mangrove, in Shorea laevifolia forest 
(Nunukan), on a dry bamboo ridge at 300 m, also 
on red or yellow sandy loams, exceptionally as 
high as 750 m, and even 1900 m. Fl. March-May 
(June-Jan.), fr. June-Aug. (Sept. -Jan.). Flowers 
are noted to be fragrant. The fruits are obviously 
buoyant, the sterile cells being filled with air. 

Vern. Sumatra: kindling, Palemb., krunjing, 
Banka; Borneo-Sarawak: purup, Lundu; Sabah: 
kayu tanyong, kulimbabok, tandjong jawa, tanjong- 
tanjong, M, mangkasugoi, Mub., songal, Tengara, 
inderatan, Bajau, balas, Banggi, enadak, inderopis, 
lamai-lamai, mala kinai, titkil-tukil, Dusun; 
Kalimantan: adad, Nunukan, bintangur pantai, E. 
Kutei, mangkinang tikus, Kahajan, tawi, Sampit. 

Notes. In Morotai I. a differing population is 
found, with tomentose axis of raceme and bracts 
and calyx lobes longer than in other specimens, and 
growing at 800 1000 m. Fig. 11. 

Also in West New Guinea (Vogelkop Peninsula) 
deviating specimens are found with large, thicker- 
walled fruits at c. 1900 m. 

15. Swnplotos terasifolia Wall. (Cat. 1831, 

n . 443"4, nomen) ex D< Prod. 8 (1844) 257; MiQ. 

Fl. Ind. Bat. I, 2 (1859) 466, excl. stlrp, /oil.: 

( i UUU . 1 I Br. Ind. 3 (1882) 580; Brand. I'll. K. 

Heft 6 (1901) 52; K. & G. J. As. Soc. Bcng. 74, ii 

J45; Ridi . l I. Mai. Pen. 2 (1923) 306; Nooi 

Leid. But. Scr. I (1975) 140, pi. 7c I Bobua 

folia Miens, J l inn. Soc. Bot. 17 (1879) J04. 

ilium O. K. Re\ Gen. PI. 2 

I iK. 7, 10c. 

| < « r.isifolia. I i^. 7, IOC 

to 2^ in, J5 an i idingly 

thin-pilose in innovati th discontinuous; 

terminal buds with many leal let, the latter 



leaving conspicuous scars. Leaves long spreadingly 
to more or less appressedly pilose beneath, espe- 
cially on midrib and nerves, sometimes entirely 
glabrous, with cuneate base, sharply dentate mar- 
gin and acuminate apex, 7-16(-22) by 2-5(-7) cm; 
nerves 6-9 pairs, meeting in a distinct looped intra- 
marginal vein; petiole slender, 15-25 mm. Spike 
resembling a short cone in bud as in S. barringtonii- 
folia, becoming at most 3 cm long; axis ± appres- 
sedly long pilose to densely pubescent. Bracts 
broadly ovoid to orbicular, boat-shaped, appres- 
sedly (silky-)pubescent on the back, at least in the 
middle, c. 5 by 5 mm; bracteoles with same indu- 
ment, narrowly elliptic, c. 3 mm long, both soon 
caducous. Calyx glabrous or slightly pubescent, 
2' 2 -4 mm, the lobes initially 1—1 Va mm. becoming 
often as long as the calyx by tearing apart. Corolla 
c. 5 mm. Stamens 30 to more than 100. Disk 
glabrous. Ovary glabrous, l-l 1 2 mm high; style 
glabrous, but the conical base sometimes hairy. 
Fruit ellipsoid, shiny blue, 22-40 by 8-18 mm; 
stone with 8 high ridges, 3-celled with a central 
canal, often only one cell developed. Seed cylindri- 
cal, with straight embryo. 

Distr. Extreme south of Peninsular Thailand; 
in Malesia: Sumatra (also Banka), Malay Penin- 
sula, Borneo, and West New Guinea (once, near 
Merauke), showing a most unusual disjunction in 
range. 

Ecol. Lowland rain-forest, hillsides on granite, 
on granitic sand, low ridges with sandy soil, also 
sandy loam with lime, mostly below 200 m, rarely 
ascending to 1000 m. Fl. June, fr. April-Oct. 

Vern. Sumatra: sisiham, Pakanbaru, minta- 
pung, mentepung, Banka. 

b. var. grandifolia Noot. Leid. Bot. Ser. 1 (1975) 
141. 

Leaves c. 30 by 8 cm. Nerves 10-14 pairs. 

Distr. Malesia: NE. Sumatra (Asahan), 2 collec- 
tions. Flowers unknown. 

16. Symplocos cochinchinensis (Lour.) S. Moore, 
J. Bot. 52 (1914) 148; Guillaumin, Bull. Soc. Bot. 
Fr. 71 (1924) 277; Fl. Gen. I.-C. 3 (1933) 998; 
MhKR. Comm. Lour. (1935) 304; Hand. -Ma//. 
Beih. Bot. Centralbl. 62 B (1943) 32; H. L. Li, J. 
Wash. Ac. Sc. 43 (1953) 107; Noot. Leid. Bot. 
Ser. I (1975) 141, with full synonymy. — Dicalix 
cochinchinensis Lour. Fl. Coch. 1 (1790) 663, 

excl. tyn. Arbor rediviva RUMPH. Fig. 12, 13. 

For the many synonyms sec under the varieties. 

Small shrub to large tree, heaves very variable in 

all characters. Inflorescence usually a spike, rately a 
raceme, but in up. leptophylla sometimes red u ced 
to .i fascicle in the axils o\ the leaves or beneath 
them, in Up. thwaltesii sometimes a panicle ol 

racemes. Fruits ampulliform to globose, in up. 
leptophylla and up. thwaltesii from globose to 

ellipsoid, OVOld oi ampulliform, in W cochinchin- 
ensis sot . imbrtcata ovoid to ellipsoid. Seed and 
embryo curved. 
Distr. (ontment.il Asia (India, Burma, rhai- 

laml, IihIo-( hma. < lima, Japan. Ryu Ksu Is . 
Hainan. I oimosa), thrOUghOUl Mulcsni to Austia 

lia (Queensland, Ne* South Wales, I ord Howe I.), 

the Solomons, New Hebrides, anil 1 i|i 

Notes. I he oldest name fbl this species is 
HyrtUt laurinui Km/. I7K(,. However, its epithet 



246 



Flora Malesiana 



[ser. I, vol. 8 2 



can not be used because of the heterotypic synonym 
S. lamina Wall, ex G. Don, 1837. 

This is the most widely distributed and also most 
variable species of the genus. The two main forms 
of the western part of its distribution, 'cochinchinen- 
sis'' and i laurina\ have usually been treated as 
different species, the main difference being hairy 
versus glabrous calyx lobes; in addition the bract 
and bracteoles in cochinchinensis form a cup 
appressed to and concealing the ovary while the 
calyx lobes often enlarge in fruit forming a conical 
beak. In lamina the cup formed by the bract and 
bracteoles is more platter-shaped, while the calyx 
lobes form a small crown on top of the fruit, but 
they can also be closed. 

These two forms can be kept rather well apart in 
large parts of the range, but in other parts they keep 
less well separate and this results in a great varia- 
bility, in part intergrading, which I have ascribed 
to hybridization, while it is possible that from these 
hybrid swarms new small local taxa may have 
evolved through environmental conditions, e.g. 
var. sessifolia and var. imbricata. 

Towards the eastern end of the range, in New 
Guinea, Australia, and the Pacific Islands forms 
occur which often have no resemblance any more 
to the two main western forms, but in the inter- 
mediate area they are linked with them in a con- 
tinuous variation, and thus break down any defin- 
able distinction between them. 

In these eastern forms, which I assume are 
'derived' during the former eastward extension of 
the range, some new tendencies have developed, in 
that seed and embryo are only curved at the base 
and are uncinate and that there is a tendency to- 
wards unisexuality of the flowers. Several New 
Guinean forms are further characterized by a 
condensed fascicle-like inflorescence, while the 
disk often becomes hairy. 

Within the species 5 of the 9 pollen subtypes 
known from subg. Hopea are found. The pollen 
type is only constant for ssp. laurina and for ssp. 
cochinchinensis and its varieties philippinensis and 
sessifolia. 

Instead of giving a lengthy discussion on the 
variability I have found it more convenient and 
clear to subdivide the species in formally named 
subspecies and varieties, although I am aware that 
it will not always be possible to name odd deviating 
or intermediary specimens. 

KEY TO THE SUBSPECIES 

1. Seeds and embryo twice curved. Inflorescence a 
basally branched spike, rarely a raceme. Flowers 
bisexual. Disk always glabrous. Fruit ampulli- 
form (ovoid to ellipsoid in ssp. cochinchinensis 
var. imbricata). 
2. Calyx lobes hairy (except on Mt Dieng in 
Central Java), often enlarged in fruit, forming 
a conical beak .... 1. ssp. cochinchinensis 
2. Calyx lobes glabrous, often ciliate, not en- 
larged in fruit 2. ssp. laurina 

1. Seeds and embryo once curved. Inflorescence a 
basally branched spike or raceme, or flowers 
solitary or in a fascicle. Flowers bisexual or 
functionally unisexual (or plant polygamous). 
Disk glabrous or hairy. Fruit ellipsoid to ovoid 
or ampulliform. 



3. Seeds and embryo once curved. (Disk glabrous 
or rarely pilose.) Calyx lobes glabrous, often 
ciliate. Flowers bisexual . . 3. ssp. thwaitesii 

3. Seeds and embryo uncinately curved towards 
the base. (Disk glabrous to densely pilose.) 
Calyx lobes glabrous to densely hairy. Flowers 
functionally unisexual or polygamous (in male 
flowers the stigma is absent) 4. ssp. leptophylla 

1. ssp. cochinchinensis. 

For synonyms see under (he varieties. 

KEY TO THE VARIETIES 

1 . Leaves usually pubescent or tomentose beneath ; 
nerves 10-14 pairs, much prominent beneath, 
strictly parallel to each other, nearly reaching the 
margin; petiole (2-)5-17(-35) mm. 

a. var. cochinchinensis 
1. Leaves glabrous; nerves 4-11 pairs, usually not 
strictly parallel to each other, anastomosing or 
meeting in an intramarginal vein at some dis- 
tance of the margin. 
2. Leaves 3-12 by lV 2 -6 cm, index 1V2-3; nerves 
4-8 pairs; petiole 0-3 mm. Fruit at most 7 mm 

long b. var. sessifolia 

2. Leaves 6-18 by l'/^ViCm, index IV2-4V2; 
nerves 5-11 pairs; petiole 3-25 mm. Fruit at 
most 7 mm long . . c. var. philippinensis 
2. Leaves 4-9 by 2 l / 2 -5 l l 2 cm, index 1-2; nerves 
5-7 pairs; petiole 4-7 mm. Fruit 10-12 mm 
long d. var. imbricata 

a. var. cochinchinensis. — Dicalix cochinchinensis 
Lour. Fl. Coch. 1 (1790) 663, excl. syn. Arbor 
rediviva Rumph. — Dicalyx aluminosus Bl. Bijdr. 
(1826) 1117, p.p. — Dicalyx javanicus Bl. I.e. 
1 1 17. — S.ferruginea Roxb. (Hort. Beng. 1814, 40; 
Wall. Cat. 1 83 1 , n. 441 2, nomen) Fl. Ind. ed. Carey 
2 (1832) 542; Miq. Fl. Ind. Bat. 1, 2 (1859) 466; 
Clarke, Fl. Br. Ind. 3 (1882) 574; K. & V. Bijdr. 7 
(1900) 141 ; Brand, Pfl. R. Heft 6 (1901) 40; K. & 
G. J. As. Soc. Beng. 74, ii (1906) 238; Koord. 
Atlas 2 (1914) t. 384; Ridl. Fl. Mai. Pen. 2 (1923) 
302. — S. mollis Wall. (Cat. 1831, n. 4433, 
nomen) ex G. Don, Gen. Syst. 4 (1837) 3. — S. 
spicata Roxb. var. platystachya G. Don, I.e. 2. — 
5. polystachya Wall. (Cat. 1831, n. 4428, nomen) 
ex DC. Prod. 8 (1844) 254; Mor. Syst. Verz. (1854) 
43; Zoll. Syst. Verz. 2 (1854) 136; Miq. Fl. Ind. 
Bat. 1, 2 (1859) 465. — S. verhuellii Jungh. & 
De Vr. PI. Ind. Or. 3 (1845) 12; Miq. Fl. Ind. Bat. 
1, 2 (1859) 467. — S. horsfieldiana Miq. Sum. (1861) 
475. — S. lachnobotrys Miq. I.e., inch var. glabrior 
Miq. — S. javanica Kurz, J. As. Soc. Beng. 40, ii 
(1871) 64; ibid. 46, ii (1877) 239, excl. syn. S. rubi- 
ginosa\ Merr. Int. Rumph. (1917) 420; Heyne, 
Nutt. PI. (1927) 1263; Burk. Diet. (1935) 2114; 
Back. & Bakh./. Fl. Java 2 (1965) 205. — Lodhra 
javanica Miers, J. Linn. Soc. Bot. 17 (1879) 302. — 
Lodhra ferruginea Miers, I.e. 299. — Lodhra 
polystachya Miers, I.e. 300. — Lodhra verhuellii 
Miers, I.e. 302. — S. ferruginea Roxb. var. 
polystachya Clarke, Fl. Br. Ind. 3 (1882) 575. — 
Eugeniodes ferrugineum O. K. Rev. Gen. PI. 2 
(1891) 975. — Eugeniodes lachnobotryum O. K. I.e. 

— S. delectans Brand, Bot. Jahrb. 54 (1916) 219. 

— 5. ferruginea Roxb. var. delectans Kaneh. & 
Hatus. Bot. Mag. Tokyo 56 (1942) 487. — Fig. 7. 



1977] 



Symplocaceae (Nooteboom) 



247 



Shrub or small tree, 9-22 m by 30 cm 0, rarely a 
large tree to 45 m by 80 cm 0. Twigs rusty tomen- 
tose or velvety, glabrescent, rarely pubescent, 
appressedly pilose, or glabrous. Leaves rusty or 
brownishly pubescent or tomentose beneath, 
especially on midrib and nerves, rarely glabrous, 
(ovate to) elliptic (to obovate) with cuneate, rarely 
rounded or (in New Guinea) cordate base, glandu- 
lar dentate or crenulate margin and more or less 
acuminate apex, (6— )12-25 by (2\' 2 -)3-10 cm; 
nerves (8— )10— 14(— 16) pairs, very prominent 
beneath, parallel to each other, mostly quite 
straight, curved upwards towards the margin and 
nearly reaching it, whether forming an intramargi- 
nal vein or not; petiole (2-)5-17mm (rarely to 
35 mm in New Guinea). Spike usually branched, 
3-15 cm, in topodeme morobeensis up to 3 cm, axis 
densely rusty tomentose or pubescent, in New 
Guinea sometimes sericeous. Bracts and bracteoles 
persistent, with same indument, the former at least 
2 mm long and broad, but usually longer, excep- 
tionally up to 10 mm long, with the 2 smaller 
bracteoles forming a calycle hiding the ovary. 
Flowers faintly scented to fragrant. Calyx appres- 
sedly pubescent (in topodeme morobeensis indument 
only towards the apex), divided into (l-)2(-3) mm 
long lobes. Corolla white (according to some collec- 
tors with a yellow spot on each lobe), from 2 
(sometimes in New Guinea) to 3-5 mm long. 
Stamens 30-70 (in New Guinea from 10 at high 
altitudes to more than 70 at low altitudes). Disk 
glabrous. Ovary glabrous, ' 2 -l mm high; style 
glabrous, 3-5 mm. Fruit ampulliform or globose, 
5-7 by 4-5 mm, more or less ribbed when dry, 
often narrowed into a cylindrical neck, crowned by 
the usually closed, enlarged, calyx lobes which form 
a conical beak on top. Seed 1, twice curved with 
similar curved embryo. 

Distr. Continental SE. Asia (India, Burma, 
Thailand, Indo-China, China, Hainan, Formosa, 
Ryu Kyu Is., Japan) and throughout Malesia 
except the Lesser Sunda Is., Celebes, and the 
Moluccas, scarce in the Philippines. 

Ecol. A variety of habitats over a considerable 
altitudinal range, from the lowland up to c. 
2500 m, in New Guinea even to 3000 m, in the 
understorey of rain-forest, primary and secondary, 
in the hills often associated with Eugenia and 
Fagaceae, extending to a few exceptional condi- 
tions, e.g. in Banka and Billiton on granite sands. 
Fl. (Jan. -May) June -Sept. (Oct. Dec.), fir. Oct.- 
July. Ripe fruit dark blue. In Malaya crown shape 
often called deep, domed, narrow and dense. 

Vern. Sumatra: dige~ra, k€dung, ke~mbang lonah, 
Djambi, kayu njari badok, Lampong, kaju salon- 
dung, k. si hondung, Padanglawas, kikaput, 
Pasemah, loba-loba, Batak, madang harbo, Tapa- 
nuli, mtnkindung, Banka, se~ke~ndum, se~ panda ng, 
Palcmbang; Java: djirak, S, ki html, Muntam; kavu 
ara, Kota Belud, habo, Sg. Baru, kayu (huihu, 
Bandjar, Martapura; Philippines: tabu, Ifiago; 
New Guinea: kumin, Wigotc, Wapi lang., kutomi, 
Wandammcn lang., mirik, Sepik, Waskuk lang. 

Notes. Var. cochinchinensia possesses rather 
constant characters in large paris <>l If 
especially in continental Asia In Java cUbftMU 
bgCffffi g rather common, r | i Java 

the number of nerves decreases, and the leaves 
begin to resemble those of ssp. luurina. Merc we 



find the gradual transition to var. philippinensis. 
The latter variety replaces var. cochinchinensis in 
the Lesser Sunda Is., Celebes, the Moluccas, and 
most of the Philippine islands. 

A conspicuous population from the Morobe 
District, New Guinea, is named topodeme moro- 
beensis (petioles 1 5-35 mm, inflorescence up to 3 cm, 
indument of calyx only towards the apex or on the 
margin). 

b. var. sessifolia (Bl.) Noot. Leid. Bot. Ser. 1 (1975) 
153. — Dicalyx sessifolius Bl. Bijdr. (1826) 1118. 
— Dicalyx salaccensis Bl. I.e. — S. laurina (non 
Wall.) Mor. Syst. Verz. (1845) 42. — S. sub- 
sessilis Choisy (ex Zoll. Syst. Verz. 2, 1854, 136, 
nomen) ex Miq. Fl. Ind. Bat. 1, 2 (1859) 467. — S. 
sessi(li)folia Gurke in E. & P. Nat. Pfl. Fam. 4, 1 
(1890) 170; Brand, Pfl. R. Heft 6 (1901) 35; 
Koord. Atlas 2 (1914) t. 388; Back. & Bakh. /. 
Fl. Java 2 (1965) 205. — Eugeniodes sessilifolius 
O. K. Rev. Gen. PI. 2 (1891) 409. — Eugeniodes 
salaccense O. K. I.e. — Eugeniodes diengense O. K. 
I.e. — S. spicata Roxb./. subsessilis K. & V. Bijdr. 
7 (1900) 146. — 5. cochinchinense ssp. sessifolia 
Noot. ex Steen. Mt. Fl. Java (1972) pi. 52-4. — 
Fig. 7, 12, 13c-e. 

Shrub 1-5 m to small tree, 10 m, 10 cm 0. 
Twigs glabrous or nearly so. Innovations purple. 
Leaves glabrous, coriaceous, with cuneate- 
attenuate base and faintly acuminate apex, 3-12 by 
1 l 2 -6 cm ; nerves 4-8 pairs, meeting in a faint 
intramarginal vein; petiole 0-3(-5) mi l. Spike 
often branched, up to 6 cm, often crowdec towards 
the end of the twigs, axis densely appressedly 
pubescent; flowers purplish. Bracts, bracteoles and 
flowers as in var. cochinchinensis, but on Mt Dieng 
the calyx only ciliate, or only pubescent towards 
the margin. Calyx lobes on the fruit not enlarged 
and closed. 

Distr. Malesia: West & Central Java (Mts Salak 
eastward to Sumbing). 

Ecol. A constituent of the summit forest of the 
volcanic peaks, often associated with Myrsine, 
Leptospermum, Eurya, Schima, Photinia, and 
Myrica, on stony ridges and summits, able to 
invade exposed sterile rocky places in the vicinity 
of craters as a dwarf pioneer shrub, 1700-3050 m. 
Fl. mainly Oct. -Jan. (Febr.-March), fr. July-Aug. 

For the ecology and flower biology see the 
general paragraphs under the genus. Fruit blue- 
black when ripe. Flush purple or blue-volet. 

Uses. Flush is sometimes eaten as lalab (veget- 
able). 

Vern. Djirak, putat, S, djirik me~lowo, sasah, J. 

c. var. philippinensis (Brand) Noot. Leid. Bot. 
Ser. 1 (1975) 154. Dicalyx aliinunosu> Bl Hi tdi 

(1826) 1117,9.9, v micata (non Roxb.) F.-Vnx. 

Nov. App. 4 (IKK0) 127. S. syrtngoldu Brand, 
Pfl, K. Heft 6(1901)41 ,s. Moore, J. Bot 52 ( i vi-;> 
148; Mirr. Int. Rumph. (1917) 421 s ft! 

ruftnea Roxi var. philippinensis Brand, Philip. J. 

| 1908) Bot, 6 S. alninii Brand, /, ; 

Mirr. I n Philip. 1 (1923) 297 V romosiiMm*. 
Philip. J. s, i. 1 (191 r) Bol 293 I a. Philip. 3 
(1923) J02, S ferruginea Roxb. var. tyringokkt 
Ham. f. Bcih Bot < entralN 19 B (1923) 
s favanka(non Ki u) Mm I n. Philip. 3(1923) 
299. Fig. 7. 



248 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 12. Symplocos cochinchinensis (Lour.) S. Moore ssp. cochinchinensis var. sessifolia (Bl.) Noot. in 
fruit (and 1 flower), on summit of Mt Pangrango, West Java, at 3000 m (Nooteboom 906). Photogr. 

Nooteboom, Febr. 1969. 



Tree to 25 m high, 50 cm 0. Twigs glabrous or 
appressedly pubescent. Leaves glabrous, or the 
midrib sparsely appressedly fine-hairy, ± elliptic 
with cuneate base and acuminate apex, (4-)6-18 by 
IV2-6V2 cm, but 5-13 by 2 1 l z -5 l j 2 cm in the Philip- 
pines; nerves 5-10(-15) pairs; petiole 3-15, in the 
Philippines 10-25 mm. Spike with tomentellous to 
pubescent axis. Bracts 1-2 mm, to 3 mm in the 
Lesser Sunda Is., with the bracteoles with same 
indument as the spike, further as in ssp. laurina. 
Calyx finely appressedly pubescent, divided into 
c. 1 mm long lobes. Corolla 3-6 mm. Stamens 
35-70. Disk glabrous. Ovary glabrous, V2-1 mm 
high; style glabrous, 2-5 mm. Fruit as in ssp. 
laurina. 

Distr. Malesia: Central & East Java, Lesser 
Sunda Is. (Bali, Sumbawa, Flores), Philippines 
(common, throughout), Celebes, Moluccas (Tidore, 
Ternate, Bum, Ambon, Ceram). 

Ecol. In Java in mountain rain-forest, also in 
tjemara forest, 700-2600 m, in the Lesser Sunda Is. 
500-2400 m, in the Philippines from low altitude 
up to 2000 m, also recorded from primary Diptero- 
carp forest, in the Moluccas from low altitude to 
1400 m. Fl. (Jan. -June) July-Dec, fr. Jan.-Aug. 
Flowers said to be scented; fruit turning through 
red to blue. 

Vern. Java: kayu djurang, tjirug, J; Philippines: 
abuabu, chaniusiu, gudik, Ig., banatong-babde, Tag., 
tarangisi, Bag., ngarau-ngarau, Neg. ; Moluccas: 
bunga ajang, Ambon. 

Note. In East Java this variety has probably 
originated by hybridization between ssp. laurina 



var. laurina and ssp. cochinchinensis var. cochin- 
chinensis. 

d. var. imbricata (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 155. — S. imbricata Brand, Philip. J. Sc. 4 
(1909) Bot. 109; ibid. 7 (1912) Bot. 31 ; Merr. En. 
Philip. 3 (1923) 299. — Fig. 7. 

Shrub or tree, 8-10 m. Twigs glabrous. Leaves 
glabrous, more or less coriaceous, usually broadly 
ovate, with cordate or slightly acuminate base, 
glandular dentate margin, and acuminate apex, 
4-9 by 2 1 /2-5 1 /2cm; nerves 5-7 pairs. Spikes axil- 
lary or pseudoterminal. Bract and bracteoles per- 
sistent. Calyx more or less appressedly pubescent, 
divided into 2 mm long lobes. Corolla c. 6 mm. 
Stamens c. 60. Disk glabrous. Ovary glabrous, 
c. 1 mm high. Fruits black, ovoid to ellipsoid, 
10-12 by c. 8 cm with smooth stone. 

Distr. Malesia: Philippines (Luzon). 

Ecol. In subalpine one-storey, mossy forest, 
2000-2600 m. Fl. Dec-April, fr. Aug.-Sept., Jan- 
March. Innovations glossy redbrown, ripe fruit 
dark blue. 

Note. This mountain form is probably directly 
derived from var. philippinensis, from which it 
differs in the shorter leaves and larger fruit. 

2. ssp. laurina (Retz.) Noot. Leid. Bot. Ser. 1 
(1876) 156. — Myrtus laurinus Retz. Obs. Bot. 4 
(1786) 26. 

Note. This subspecies ranges from Ceylon 
eastwards to Celebes, China and Japan. Besides 
the type variety there is only one local stenophyl- 
lous variety in Indo-China and S. China. 



1977] 



Symplocaceae (Nooteboom) 



249 




Fig. 13. Leaf sue and shape in Symplocos cochin- 
ii mi i i s Mooiu at different altitudes, 
from a to c at 100, 1600, 71800, 2000, and KKK) in 
respectively, a b. wp. laurina (Km/.) N<xm. var. 
kmrina, c e. up. cochlnchineruii vat teutfblla (Bl.) 
Noot., all • ,<,i i i uihai'ms 4561,6 Blokhi is 
7 12 21, c Blum 1965, rfJa 4010, s-Da raw van 

I n i win 8425). 



e. var. laurina. — Law us sen at a floris spicatis 
Burm. Thes. Zeyl. (1737) 139, t. 62. — Myrtus 
laurinus Retz. Obs. Bot. 4 (1786) 26, non S. laurina 
Wall, ex G. Don, 1837. — Drupatris cochinchin- 
ensis Lour. Fl. Coch. 1 (1790) 314. — Decadia 
aluminosa Lour. I.e. 315. — Eugenia laurina 
Willd. Sp. PI. 2 (1799) 967, p.p. — Dicalyx 
spicatus Bl. Bijdr. (1826) 1118. — Dicalyx acumi- 
nata Bl. I.e. 1119. — 5. jp/Va/a Roxb. (Hort. 
Beng. 1814, 40; Wall. Cat. 1831, n. 4417, nomeri) 
Fl. Ind. ed. Carey 2 (1832) 542; Choisy in Zoll. 
Syst. Verz. 2 (1854) 136; Miq. Fl. Ind. Bat. 1, 2 
(1859) 465; Clarke, Fl. Br. Ind. 3 (1882) 573, 
incl. var. malasica I.e. et var. laurina I.e. p.p. ; K. & 
V. Bijdr. 7 (1900) 144, incl. f. javanica I.e. et f. 
acuminata et f. xanthophylla I.e. 145, excl. f. sub- 
sessilis; Brand, Pfl. R. Heft 6 (1901) 39, incl. var. 
acuminata I.e. 41 ; K. & G. J. As. Soc. Beng. 74, ii 
(1906) 236; Koord. Atlas 2 (1914) t. 386, 387; 
Ridl. Fl. Mai. Pen. 2 (1923) 301; S. Moore, J. 
Bot. 63 (1925) Suppl. 65; Heyne, Nutt. PI. (1927) 
1263; Burk. Diet. (1935) 2115; Back. & Bakh. 
/. Fl. Java 2 (1965) 205. — S. laurina Wall. (Cat. 
1831, n. 4416, nomeri) ex G. Don, Gen. Syst. 4 
(1837) 3; Rehd. & Wils. in Sargent, PI. Wils. 2 
(1916) 594; Rehd. J. Am. Arb. 15 (1934) 298; 
Merr. Comm. Lour. (1935) 303; Corner, Ways. 
Trees (1940) 623 ; Hand.-Mazz. Beih. Bot. Central- 
bl. 62 B (1943) 33; Steen. Fl. Mai. I, 5 (1957) 
clxxxi, f. 4. — S. polycarpa Wall. (Cat. 1831, 
n. 4423, nomeri) ex G. Don, Gen. Syst. 4 (1837) 3; 
Miq. Fl. Ind. Bat. 1, 2 (1859) 465. — S. ribes 
Jungh. & de Vr. PI. Ind. Or. 3 (1845) 1 1 ; Miq. Fl. 
Ind. Bat. 1, 2 (1859) 468. — S. acuminata Miq. 
I.e. 467. — Lodhra ribes Miers, J. Linn. Soc. Bot. 
17 (1879) 302. — Lodhra xanthophylla Miers, 
I.e. — S.ftavida Miq. (PI. Hohenacker n. 1053) ex 
Clarke, Fl. Br. Ind. 3 (1882) 573, in syn. — 
Eugeniodes ribes O. K. Rev. Gen. PI. 2 (1891) 976. 
— Fig. 7, 13a-b. 

Shrub, 3 m, to tree, 6-14 m by 30 cm 0. Twigs 
and leaves glabrous, except sometimes the very 
youngest parts. Leaves ± elliptic with cuneate base 
and acuminate apex, 4'/ 2 -21 by (I 1 /?-) 27 2 -8 cm; 
nerves 6-9 pairs (but in forms transitional to var. 
cochinchinensis up to 13 pairs), not strictly 
parallel, anastomosing at some distance of the 
margin, often meeting in an intramarginal vein; 
petiole (5-)10-15(-20) mm (in transitional forms 
the leaves arc like those of var. cochinchinensis 
except for the indument). Spike l'/j-Hcm, axis 
glabrous to more or less appressedly puberulous or 
pubescent. Bracts and bracteoles persistent, at 
most 2 mm long and broad, but usually only 1 mm, 
only enveloping the base of the ovary. A short 
pedicel exceptionally present. Calyx glabrous oi 
nearly so, divided into I 2 mm long, often ciliate 
lobes, not elongating in fruit, whether 01 not 
closed after anlhesis. Rest of QOWQI and fruit as in 

ochtnchlnensts. 
D is tr. Continental Asia (India, Ceylon, Mm ma, 
Thailand. Indo-China, < bina, Hainan, Formosa, 
Japan); m Maleaia. Sumatra (also Enggano [.), 
Mai. i> Peninsula (rare), Java (very common), 
Borneo date), ( elebei (rare), 

I Substage lice in lam-lorest, sometimes in 

potation, neai waterfall. In Malaya 
found also in sandy, tidal gtUvn < Melaleuca) forest, 

in continental Asia, Sumalia and ( eleltes liom low 



250 



Flora Malesiana 



[ser. I, vol. 8 2 



altitude to c. 2000 or 3000 m, in Java only above 
c. 1000 m. Fl. Sept.-April, fr. Febr.-Sept. 
Flowers are said to be slightly foetid to strongly 
smelling, opening early in the morning. Fruit turns 
black via blue. 

Vern. Sumatra: kayu djari manuk, Batak, 
dadak putih, diera, Enggano; Java: djirak, d. sasak, 
S, djirek, J. 

3. ssp. thwaitesii (F.v.M.) Noot. Leid. Bot. Ser. 1 
(1975) 159, with full synonymy. — S. thwaitesii 
F.v.M. Fragm. 3 (1862) 22. 

Distr. This subspecies consists of 4 varieties 
which occur in Queensland, New South Wales, and 
Lord Howe I. One of these is also found in New 
Guinea. 

Note. There is one sheet (LAE 54751) which is 
not identified to a variety; it might belong to the 
Queensland var. montana (C. T. White) Noot. 

f. var. stawellii (F.v.M.) Noot. Leid. Bot. Ser. 1 
(1975) 161. — S. stawellii F.v.M. Fragm. 5 (1865) 
60; Brand, Pfl. R. Heft 6 (1901) 37, excl. var. — 
S. spicata Roxb. var. australis Bth. Fl. Austr. 4 
(1869) 292. — Fig. 7. 

Tree up to 30 m high, 80 cm 0. Twigs glabrous. 
Leaves glabrous, elliptic with broadly cuneate base 
and not or faintly acuminate apex, 6-16 by 2V2-IO 
cm; nerves 8—1 1 pairs; petiole 5-25 mm. Spike 
often branched, rarely exceeding 6 cm, the axis 
glabrous or appressedly puberulous. Bracts and 
bracteoles usually persistent, 1— 1 1 / 2 and V2-I m m 
long respectively. Calyx glabrous, divided into 
V4- 3 /4 mm long lobes. Corolla 3-5 mm. Stamens 
25-50. Disk glabrous. Ovary glabrous, l-l'^mm 
high; style glabrous. Fruit ellipsoid-ovoid, 5-7 mm. 

Distr. Australia: Queensland, New South 
Wales, and Lord Howe I. ; in Malesia: New Guinea 
(Papua). 

Ecol. Two habitats are recorded, viz in the 
lowlands with influence of a dry season, on edge of 
savannah forest, and on the Oriomo R. in associa- 
tion with Acacia, and in the middle mountains at 
c. 2000-2300 m, in secondary forest, tall mixed 
rain-forest, and in Podocarp-dominated forest on 
peaty soil. Fl. June, Sept.,/r. June-Oct. Flowers 
are recorded to be fragrant. Fruit develops from 
green via blue to purple-black. 

Vern. New Guinea: tuliper, Poio, Enga lang., 
kurfgum, Yogoo, Enga lang., truom, Oriomo R., 
Kiunga lang. 

4. ssp. leptophylla (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 162. — S. stawellii F.v.M. var. leptophylla 
Brand, Pfl. R. Heft 6 (1901) 37. — 5. leptophylla 
Turrill, J. Linn. Soc. Bot. 43 (191 5) 30. — S. mam- 
beramo Brand, Nova Guinea 14 (1924) 186. 

For further synonyms see under the varieties. 

Notes. This is a rather heterogeneous subspecies 
ranging from the Lesser Sunda Is. and Moluccas 
through New Guinea (incl. Bismarcks) to Mela- 
nesia (Solomons, New Hebrides) and W. Poly- 
nesia (Fiji), the type having been described from 
Fiji. The varieties are rather reticulately allied and 
are often connected by intermediate specimens 
among which may be some hybrids. Some collec- 
tions I could not refer to a variety, in part due to 
inadequate material, e.g. the type of 5. mamberamo. 

In most varieties the flowers are functionally 



unisexual or bisexual in the same variety. In the 
functionally female flowers the number of stamens 
is low, while the style is large, with peltate stigma. 
In the functionally male flowers the number of 
stamens is high and the style is small, without 
stigma. 

KEY TO THE VARIETIES 

1 . Underside of leaves hairy. 
2. Twigs glabrous .... g. var. leptophylla 
2. Twigs hairy. 
3. Calyx and ovary glabrous. 
4. Disk hairy g. var. leptophylla 

4. Disk glabrous. Twigs sericeous or tomen- 
tose s. var. ovata 

3. Calyx and/or ovary hairy. 

5. Ovary glabrous. 

6. Disk glabrous. Twigs sericeous or tomen- 

tose s. var. ovata 

6. Disk hairy. 
7. Twigs (appressedly) pubescent. 

g. var. leptophylla 
7. Twigs tomentose or pilose. 

v. var. versteegii 
5. Ovary hairy. 
8. Calyx glabrous. 
9. Twigs sericeous or tomentose. 

s. var. ovata 

9. Twigs (appressedly) pubescent. 

g. var. leptophylla 
8. Calyx hairy. 

10. Disk glabrous. 

1 1 . Bracts shorter than 3 mm, fruits to 
c. 10 mm long s. var. ovata 

1 1 . Bracts longer than 3 mm, fruits more 
than 10 mm long . . t. var. revoluta 

10. Disk hairy. 

12. Twigs (appressedly) pubescent. 

g. var. leptophylla 
12. Twigs not appressedly pubescent or 
puberulous, e.g. tomentose. 
13. Petiole more than 20. 

j. var. tomentosa 
13. Petiole less than 20 mm. 
14. Inflorescence a (basally branched) 
spike. 
15. Bracts shorter than 3 mm 

o. var. reginae 

15. Bracts longer than 3 mm 

t. var. revoluta 
14. Inflorescence not a spike. 

16. Nerves in 7-11 pairs. 

m. var. molobros 
16. Nerves in 4-8 pairs. 

o. var. reginae 
1 . Underside of leaves glabrous. 
17. Calyx and ovary glabrous. 
18. Twigs hairy. 
19. Petiole to 5 mm. 
20. Leaves shorter than 5 cm. 

r. var. orbicularis 

20. Leaves longer than 5 cm. 

I. var. longilobata 
19. Petiole more than 5 mm. 

21. Leaves obovate, 10-25 cm. Petiole 10- 
40 mm i. var. insularis 

21. Leaves ovate or elliptic, 2V2-23 cm. 
Petiole 5-25 mm. 



1977] 



Symplocaceae (Nooteboom) 



251 



22. Leaves ovate or elliptic, 2 1 2 -11 cm. 
Twigs sparsely appressedly pilose. 

u. var. sogeriensis 
22. Leaves ± elliptic, 5-23 cm. Twigs 
appressedly pubescent, g. var. leptophylla 
18. Twigs glabrous. 

23. Inflorescence a very slender, often branched 
spike (or raceme) of 2-10 cm. 
24. Twigs (exceptionally) thick. 

p. var. schumanniana 
24. Twigs not (exceptionally) thick. 
25. Intramarginal vein far from margin. 

p. var. schumanniana 
25. Intramarginal vein close to margin. 

w. var. maculata 
23. Inflorescence a fascicle or a (reduced) often 

branched, stout spike (or raceme). 
26. Petiole to 5 mm . . r. var. orbicularis 
26. Petiole more than 5 mm. 
27. Bracts and bracteoles caducous. New 
Hebrides. 

var. aneityensis (Brand) Noot. 
27. Bracts persistent. 
28. Reticulation fine, usually prominent on 
both under and upper surface. 

u. var. sogeriensis 
28. Reticulation fine or coarse, usually only 

prominent on the undersurface. 
29. Leaves usually less than 5(-8) cm long. 
h. var. monticola 
29. Leaves usually more than 5 cm long. 
30. Inflorescence axis glabrous. 

q. var. floresana 
30. Inflorescence axis hairy. 
31. Leaves obovate . . i. var. insularis 
31. Leaves elliptic or circular. 

g. var. leptophylla 
17. Calyx and or ovary hairy. 
32. Petiole to 5 mm . . . x. var. parvifolia 
32. Petiole more than 5 mm. 
33. Ovary glabrous. 
34. Disk glabrous. Twigs glabrous. Reticula- 
tion fine, usually prominent on both under 
and upper surface, calyx lobes to c. 1 I 1 mm 

long u. var. sogeriensis 

34. Disk hairy. 
35. Leaves usually less than 5(-8) cm long. 

h. var. monticola 
35. Leaves usually more than 5 cm long. 
36. Inflorescence axis glabrous. 

k. var. doormanensis 
36. Inflorescence axis hairy. 

g. var. leptophylla 
33. Ovary hairy. 
37. Twigs hairy. 
38. Calyx glabrous . . v.- W. leptophylla 
38. Calyx hairy. 
39. Calyx symmetrically cleft. 

n. var. pediccllata 
39. Calyx regular. 

40. Leaves obovate ... I. var. insularis 
40. Leaves elliptic or circular. 

U- var. Ii'plophylla 
37. Twigs glabrous. 
41. Calyx glabrous v.. var. loptoplnlla 

41. Calyx hairy. 
42. Inflorescence a very slender 

branched spike (in raceme) oi ■- 10cm 

p. var. s« hum. imiii. in. i 



42. Inflorescence a fascicle or a (reduced), 

often branched, stout spike (or raceme). 

43. Calyx 2-4-lobed or symmetrically cleft, 

calyx lobes becoming longer by 

tearing n. var. pedicellata 

43. Calyx regularly 5-lobed. 
44. Leaves obovate . . i. var. insularis 
44. Leaves elliptic or circular. 

g. var. leptophylla 

g. var. leptophylla. — S. stawellii F.v.M. var. 
leptophylla Brand, Pfl. R. Heft 6 (1901) 37. — 
S. leptophylla Turrill, J. Linn. Soc. Bot. 43 (1915) 
30, incl. f. compacta Turrill, I.e. 31. — 5. pal- 
mar urn Brand, Bot. Jahrb. 54 (1916) 220. — S. 
trifurceps Brand, Nova Guinea 14 (1924) 186. — 
5. romeri Brand, I.e. — S. aggregata White & 
Francis, Proc. R. Soc. Queensl. 38 (1927) 256, 
t. 17. — S. luteifolia Kaneh. & Hatus. Bot. Mag. 
Tokyo 56 (1942) 487. — 5. turrilliana A. C. Smith, 
J. Arn. Arb. 33 (1952) 111. — Fig. 7. 

Shrub 2-3 m to tree 20-28 m by 20-45 cm 0. 
Twigs glabrous or pubescent. Leaves glabrous or 
pubescent to finely appressedly pilose beneath, ± 
elliptic, with cuneate to cordate base, entire to 
dentate margin and acuminate apex, 5-23 by 2-12 
cm; nerves 6-12 pairs, meeting in an intramarginal 
vein; petiole 5-25 mm. Inflorescence a fascicle or a 
reduced, branched spike, sometimes a spike or 
raceme to 5 cm, axis appressedly puberulous to 
pubescent or sericeous. Bracts and bracteoles 
persistent, with same indument, 1-10 and 1-4 mm 
long respectively. Pedicels 0-2 mm. Flowers 6\ ?, or 
5. Calyx 3 4 -3 mm, either entirely divided into the 
hairy or glabrous lobes or not. Corolla 2-5 mm. 
Stamens c. 10 to more than 100, in 9 flowers less 
than 20. Disk softly hairy. Ovary glabrous or 
pubescent to sericeous, 1-2'ljmm high; style 
glabrous or with few hairs towards the base, small, 
without stigma in functionally cJ flowers, with 
peltate stigma in V and 5 flowers. Fruit glabrous or 
sparsely pubescent, sessile in a fascicle or infruc- 
tescence up to 5 cm or even more, ovoid to ellipsoid 
or ampulliform, often globose, 6-15 by 4-9 mm. 

D i s t r. W. Polynesia ( Fiji ), Melanesia (Solomons 
and Santa Cruz Is.); in Malesia: Moluccas (Buru, 
Ambon, Ceram) and very common in New Guinea 
(incl. Jappen, Normanby, and Goodenough 
Is.) and the Bismarck Archipelago (New Britain, 
New Ireland). 

Ecol. Very variable, rare in the lowland, mostly 
from 900-3360 m (Ml Olio), in the lain. 
ceous forest, transition of OOtdfer-CastanopsiS' 
Nothofagui forest to grassland, moss) forest on 
ridge tops, in forest relicts o\ Quercus-Dactrydium 
forest (Arlak), once noted as a dominant on upper 
ridges, in association with Podocarpus pilgtri in 
Britain, and in Casuatina forest there // 
J. in Aug., /' Ian Dec I lowers said to he Ira 

grant I nut dark blue to purple black when mature. 

Vein. New Guinea: aibih, Mum. arilth, Non- 

dugl, keUktndt, Mi Ambua, k<>kn, Telefomin, 

Nah lang., guguma, kongumu, kimgumu, Mt Hagen, 

w.uiki lang., lellcop, Waria, matala, Mt lalawe, 
Britain, navako t New Britain, patwtedledle, 
Tari, Hull lang., pelwadUi, Mt Ne, Habono, 
punrall, Wabag, tufffaro, ypap, I ngs lang., imnn, 
Saidor, utu-utu, < ycloop Mt, Ormu lang., wapt, 
Senile, Wagu lang. 



252 



Flora Malesiana 



[ser. I, vol. 8 2 



h. var. monticola Noot. Leid. Bot. Ser. 1 (1975) 166. 

- Fig. 7. 

Shrub 2 m to tree to 16 m by 22 cm 0. Twigs 
glabrous. Leaves glabrous, ± elliptic, with cuneate 
base, entire or denticulate margin and acute or 
rounded apex, 2-8 by 1-3 cm; nerves 5-7 pairs, 
meeting in an intramarginal vein ; petiole 4-10 mm. 
Spike to lV^cm long, axis glabrous or sparsely 
appressedly hairy. Bracts and bracteoles persistent, 
glabrous, 1-2 and l-l'^mm long respectively. 
Flowers functionally unisexual or bisexual as in 
var. leptophylla. Calyx appressedly pubescent or 
puberulous to glabrous, usually divided into 1—1 1 / 2 
mm long, often purple-tinged lobes. Corolla 1-272 
mm. Stamens 15-35. Disk hairy. Ovary glabrous, 
l-lV^mni high; style glabrous. Fruit ovoid to 
ellipsoid, 8-10 by 4-6 mm. 

Distr. Malesia: East New Guinea. 

Ecol. Substage tree in mossy forest and secon- 
dary forest with much climbing bamboo, 2700- 
3500 m. Fl. April-Sept., fr. July-Aug. 

Vern. Ped-ped, Giluwe, Mendi lang. 

i. var. insularis Noot. Leid. Bot. Ser. 1 (1975) 167. 

— Fig. 7. 

Tree up to 15 m by 25 cm 0. Twigs glabrous or 
appressedly pubescent. Leaves glabrous, mostly 
broadly ovate with attenuate base and acuminate 
apex, 10-25 by 572-15 cm; nerves 6-8 pairs; 
petiole 10-40 mm. Flowers not seen. Infructes- 
cence a fascicle or spike to 5V2 cm long; fruit 
sparsely pubescent, ovoid to globose, 8—13 mm 
long. 

Distr. Malesia: East New Guinea (Louisiades: 
Sudest, Rossel & Misima Is.). 

Ecol. Substage of rain-forest, along stream- 
bank, also on a summit where dwarfed to 172 m 
tall shrub; from the lowland to 800 m. Fr. July- 
Oct. Ripe fruits black. 

j. var. tomentosa Noot. Leid. Bot. Ser. 1 (1975) 167. 

Tree to 20 m. Twigs and midrib tomentose 
beneath. Leaves mostly obovate, pubescent be- 
neath, with cuneate base and rather abruptly 
acuminate apex, 18-23 by 10-12 cm; nerves 8-9 
pairs; petiole 2-3 cm. Fascicle in the axils of the 
leaves or often beneath them, including the broadly 
boat-shaped 5 mm long bracts, the 3 mm long 
bracteoles and the calyx appressedly pubescent. 
Flowers unisexual or bisexual as in var. leptophylla. 
Calyx divided into 1-2 mm long lobes. Corolla 
4-6 mm. Stamens 25 to more than 100. Disk softly 
pilose. Ovary pubescent, 1-2 mm high. Fruit not 
seen. 

Distr. Malesia: East New Guinea (Fergusson 
L). 

Ecol. Montane rain-forest dominated by oaks, 
in the substage, 700-900 m. Fl. June. Flowers said 
to be very fragrant rose-scented. 

k. var. doormanensis (Brand) Noot. Leid. Bot. 
Ser. 1 (1975) 168. — S. doormanensis Brand, Nova 
Guinea 14 (1924) 187. — 5. dalmannensis Kaneh. 
& Hatus. Bot Mag. Tokyo 56 (1942) 487. 

Shrub or small tree, P/2 rn. Twigs sparsely pilose 
to glabrous. Leaves glabrous, coriaceous, elliptic, 
with cuneate base, entire to glandular denticulate 
margin and not or faintly acuminate apex, 6-12 by 
2V 2 -6cm; nerves 5-10 pairs; petiole 7-10 mm. 



Fascicles in the axils of the leaves or on wood, in- 
cluding the 5 mm long broadly boat-shaped bracts 
and the 3-4 mm long bracteoles appressedly (long) 
pilose to pubescent; bracts and bracteoles per- 
sistent. Flowers unisexual or bisexual as in var. 
leptophylla. Calyx appressedly pilose to pubes- 
cent, 2-3 mm long, divided into the lobes. Corolla 
4-5 mm. Stamens 30-50. Disk softly pilose. Ovary 
glabrous, lV 2 -2mm high; style glabrous. Fruit 
(immature) ellipsoid. 

Distr. Malesia: New Guinea. 

Ecol. Montane rain-forest, also in mossy forest, 
1800-2700 m. Fl. Jan.,//-. Oct.-Nov. 

I. var. longilobata Noot. Leid. Bot. Ser. 1 (1975) 
169. — Fig. 7. 

Shrub or small tree, '/ 4 -8 m by 15 cm 0. Twigs 
sparsely appressedly fine-pilose, glabrescent. 
Leaves glabrous, elliptic (to orbicular) with 
rounded to more often cuneate base, crenulate 
margin and acute (to rounded) apex, 10-23 by 
6-14 mm; nerves 2-4 pairs; petiole 2-3 mm. 
Flowers unisexual or bisexual, solitary or c. 3 in a 
condensed spike to 1 cm, axis pubescent. Bracts 
and bracteoles persistent, 3-5-6 together, narrowly 
triangular, 3-5 mm long. Calyx glabrous, 272- 
4V 2 mm long, the lobes (ovate to) triangular, ciliate, 
glandular, 2-4 mm. Corolla 3-4 mm. Stamens 
14-24. Disk shortly pubescent. Ovary glabrous, 
1-2 mm high, style glabrous, 2-4 mm. Fruits ovoid 
to ellipsoid, c. 10 by 6 mm, stone rather smooth. 

Distr. Malesia: East New Guinea (Mt Wilhelm). 

Ecol. Alpine shrubberies and forest edges, in 
subalpine tussock grassland, along creek in peaty 
grassland, a stiff, fastigiate, microphyllous race, in 
sterile exposed places often dwarfed, 3200-3400 m. 
Fl. June-July, fr. July. Ripe fruit blue-black. 

m. var. molobros (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 169. — S. molobros Brand, Bot. Jahrb. 54 
(1916)217. — Fig. 7. 

Small shrub l l 2 -\ l j A m to slender tree, 4-6 m. 
Twigs densely (woolly) pilose. Leaves softly pilose 
beneath, (broadly) elliptic, with cuneate to rounded 
or even subcordate base, entire to glandular den- 
tate margin and apex whether or not acuminate, 
6-18 by 3V2-8 cm; nerves 7-11 pairs, meeting in a 
looped intramarginal vein; petiole 5-10 mm. 
Inflorescence a much reduced, branched, spike or a 
fascicle in the axils of the leaves or on wood, up to 
2 cm long; axis rusty patently sericeous-pilose. 
Bracts and bracteoles persistent, rusty long pilose 
to appressedly sericeous, 2-4 and 1-3 mm respec- 
tively. Calyx appressedly rusty sericeous or long 
pubescent, divided into 1-2 mm long lobes. Corolla 
272-5 mm. Stamens 20-60. Disk pilose. Ovary 
greyish sericeous, 1-2 mm high; style glabrous. 
Fruit ovoid to globose, 10-15 mm long, pubescent, 
becoming glabrous. 

Distr. Malesia: New Guinea. 

Ecol. Substage treelet in montane rain-forest, on 
sandy clay, on limestone or sandstone ridges, 700- 
2200 m. Fl. April-Nov., //-. Sept. 

Vern. Chandujant, Wabag, Enga lang. 

n. var. pedicellata Noot. Leid. Bot. Ser. 1 (1975) 
170. — Fig. 7. 

Shrub 2-47 2 rn to slender tree 8-16 m. Twigs 
glabrous. Leaves glabrous, stiff, ± elliptic, with 



1977] 



Symplocaceae (Nooteboom) 



253 



cuneate to rounded base and (abruptly) acuminate 
apex, 5-11 by 2 l 2 -6cm; nerves 6-10 pairs; 
petiole 5-16 mm. Raceme up to 4 cm; axis sparsely 
appressedly puberulous as the persistent 1-2 mm 
long bracts and the 1-3 mm long pedicel. Calyx 
appressedly puberulous, c. 2 mm long, wholly 
symmetrically cleft. Corolla 3-4 mm. Stamens 
c. 40 in c? flowers, c. 10 in $ flowers. Disk softly 
pilose. Ovary appressedly puberulous, 2 mm high ; 
style c. 3 mm, with conical pubescent base. Fruit 
ovoid to ampulliform, 10-15 by 7-9 mm. Seed 
strongly ruminate, embryo probably curved. 

Distr. Malesia: East New Guinea. 

Ecol. Substage of mossy forest and subalpine 
forest dominated by Nothofagus-Weinmannia or 
conifers (Araucaria, Podocarpus, Papuacedrus), 
sometimes abundant on ridges, also on limestone, 
2100-2900 m. Fl. (Jan.) April-Oct., fr. June. 

Vern. Ypap, Wabag, Enga lang., keh, kepilam, 
Enga lang. 

o. var. reginae (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 171. — S. reginae Brand, Bot. Jahrb. 54 
(1916)214. — Fig. 7. 

Shrub 1-2 m to small tree to 10 m by 10 cm 0. 
Twigs densely short and long pilose, only long- 
pilose, or woolly to tomentose; growth discon- 
tinuous. Leaves pubescent beneath, especially on 
the nerves, elliptic, with acuminate to rounded 
base, entire to glandular denticulate margin and 
acuminate apex, l' 2 -ll by 3 ,4-6 1 / 2 cm; nerves 
4-8 pairs; petiole 2-10 mm. Flowers solitary or 
few together in the axils of the leaves or below 
them, or on the apical part of an up to 3(-7) cm 
long spike; axis patently pilose. Bracts and brac- 
teoles persistent, appressedly pilose, 2-4 mm and 
1-2 mm respectively. Calyx appressedly pilose, 
divided into 1-1 ' 2 mm long lobes. Corolla 2-3 mm. 
Stamens 10-25. Disk pilose. Ovary appressedly 
pilose, 3 4 -2 mm high ; style glabrous or with pilose 
base. Fruit ovoid, pubescent, 9-15 by 7-8 mm. Seed 
1-2, curved towards the base. 

Distr. Malesia: New Guinea. 

Ecol. Oak and beech forest, also on ridges, and 
in river gorge, 900-2000 m. FL June-Aug., fr. 
Jan. -Oct. Fruit from cream through purple to 
purplish-blue when ripe. 

Vern. Dorso, Kassam Pass, Kainantu, mongutl, 
Hagen, harkomerinkey, Okapa, marnele, Morobe, 
Wagau. 

p. var. schumanniana (Brand) Noot. Leid. Bot. 
Ser. I (1975) 171. — .V. rhynchocarpa K.S< n. ex 
Brand in K.Sch. & Laut. Nachtr. (1905) 347; Bot. 
Jahrb. 54 (1916) 223. — S. schumanniana BRAND, 

17 el 224. - .V. schlechteri Brand, /.. 
224. - S. rupcstns Brand, Bot. Jahrb. 54 (1916) 
220. — S. myrmecophila S< in in ex Brand, I.e. 
224.— S.pusllllfloraS. Moore, [Vans. I inn. Sol 
II, Bot. ') (1916) 107. S. cyclopi Mrand, Nova 
Guinea 14 (l'J24) 1 88. .V. lamu BRAND, I.e. 

Pta.7. 

Shrub 2 m to tree 10 18 m by 12 37 a 
Twigs sometimes very thick, glabrous, sometimes 
iiions appressedly pubescent, often the 
branches thickened m some places, hollow, lodging 

ants Leaves • elliptic, glabrous, with ct 

entire margin end acumii ' 13 b) 

V, 14cm;ncrves8 I 5 pairs, meeting m mtramar- 



ginal vein far from the margin; petiole 5-22 mm. 
Inflorescence a slender spike (or rarely a raceme) to 
6 cm, often branched towards the base, rarely for 
its whole length; axis pubescent or puberulous to 
glabrous. Bracts and bracteoles mostly persistent, 
rarely caducous, pubescent or puberulous, l-2'/ 2 
mm and V'j-l 1 ^ mm long respectively. Pedicel if 
present at most 1 mm. Calyx glabrous or puberu- 
lous, entirely divided into c. l /a rnm long lobes, or 
l 1 2 mm long and then the lobes c. 1 mm. Corolla 
l 1 2 -5 mm. Stamens 10-30 in ? and $ flowers, 
30-80 in d and 2 flowers. Disk pilose. Ovary glab- 
rous or puberulous, 1-1 l l 2 mm high; style glabrous 
or with some hairs towards the base. Fruit ampulli- 
form, 5-6 by 3-4 mm, sometimes with rather long 
neck; stone ampulliform, rather smooth. Seed 1, 
curved, U-shaped with U-shaped embryo. 

Distr. Malesia: Moluccas (Morotai), New 
Guinea, New Ireland, and New Britain. 

Ecol. In high lowland rain-forest, sometimes 
with climbing bamboo, montane rain-forest on 
ridges, also on sandy clay, in Nothofagus dominated 
rain-forest on peaty soil, in New Britain also on 
limestone, from sea-level to 2100(-2820) m. Fl. 
Jan. -Dec, fr. July-Nov. Flowers are said to be 
faintly fragrant. Fruits turn from green through red 
to bluish when mature. 

Vern. Moluccas: reha, Morotai; New Guinea: 
pai, Wandammen, tembek, Telefomin. 

q. var. floresana Noot. Leid. Bot. Ser. 1 (1975) 172. 

— Fig. 7. 

Small, glabrous tree, up to 7 m by 15 cm 0. 
Leaves (broadly) elliptic with cuneate to rounded 
base and not or slightly acuminate apex, 9-16 by 
5-10 cm; nerves 7-12 pairs, meeting in an intra- 
marginal vein; petiole stout, 2 l l 1 -4 l l 2 cm. Spike 
basally branched, to 7 cm, axis glabrous. Bracts 
and bracteoles persistent, glabrous or appressedly 
pubescent, often ciliate. Calyx glabrous, divided 
into c. 1 mm long lobes. Corolla 3-4 mm. Stamens 
25-35. Disk glabrous. Ovary glabrous, ^^-^U mm 
high ; style glabrous. Fruit c. ovoid, 5-6 by 4-5 mm. 

Distr. Malesia: Lesser Sunda Is. (Flores). 

Ecol. Montane rain-forest, 1000-1500 m. Fl. 
May-July, fr. April. Ripe fruit blue. 

r. var. orbicularis (Hemsl.) Noot. Leid. Bot. Ser. 
1 (1975) 173. — S. orbicularis Hemsl. Kew Bull. 
(1899) 105. — S. englishii Hemsl. I.e. — S. klossii 
S. Moore, Trans. Linn. Soc. II, Bot. 9 (1916) 108. 

- Fig. 7. 

Still, often compact, microphyllous trcclct. with 
densely foliaged twigs and patent, brittle, thick 
(living fleshy) leaves; 20 50 cm to 3 10 m by 
35 cm 0. Twigs glabrous or hairy. I eaves glabrous, 

Orbicular tO elliptic, with cuneate tO rounded or 
slightly cordate base, dentate to denticulate margin 

and rounded or acute apex, 1 . i{ V >hy' .. 2 cm; 

nerves 2 7 pans; petiole I 3 mm. I lowers solitary 
Or in S spike tO 4 cm ; bracts I 3 mm, several when 

flowers solitary, oi I, Bracteoles mostly persistent, 
glabrous oi hairy, ' ■ J mm long ( sly* glabrous, 
entirel) divided into i r , nun lone lobes oi s tube 
ol 'i mm present < ©rolls 6) mm. 

Stamens liom less than 10 in , floweTS tO 25 in I 

and , flowi Di Ovarj glabrous, 

c (1-2 mm high. I ruil ellipsoid, i 15b) 4 6mm 
Disti Malesia: New Guinea. 



254 



Flora Malesiana 



[ser. I, vol. 8 2 



Ecol. Subalpine grassland shrubberies (often 
ericoid), sparse ridge top scrub, in moss-mounds 
in ridge thickets, associated with Eurya, Dimorphan- 
thera, Drimys, on creviced faces and ridges of 
sandstone, also in subalpine moss forest, bank of 
a mountain torrent, still recorded as a tree of 10 m 
at 3300 m, 2500-3800 m, in Arfak as low as 1900 m. 
Fl. June-Aug.,/r. June-Sept. 

Vern. Dibenkur, Chimbu, pombor, Giluwe, 
Mendi lang. 

s. var. ovata Noot. Leid. Bot. Ser. 1 (1975) 173. — 
Fig. 7. 

Shrub 3 / 4 -4 m to tree 12-21 m by 15 cm 0. 
Twigs appressedly sericeous to pubescent or tomen- 
tose, glabrescent, rarely glabrous. Leaves appres- 
sedly thin-hairy underneath, ovate to elliptic, with 
cuneate to cordate base and acuminate apex, 4-12 
by 2-7 cm; nerves 5-10 pairs; petiole 5-20 mm. 
Spike basally branched, axis finely pubescent to 
tomentose. Bracts and bracteoles persistent, with 
same indument as axis or less hairy, 1-3 and 1-2 
mm long respectively. Calyx glabrous but ciliate, 
or appressedly fine-hairy, divided into l lr-^ 1 li(-2) 
mm long lobes. Corolla 2-3(-4) mm. Stamens 
8-25. Disk glabrous. Ovary glabrous or sparsely 
appressedly fine-hairy, l-lV2(-2)mm high; style 
glabrous. Fruit ellipsoid to ovoid, 5-10 by 3-8 mm ; 
stone ovoid, rather smooth. Seeds 1-2, ruminate, 
fitting into the grooves of the stone. 

Distr. Malesia: East New Guinea, very 
common. 

Ecol. Substage tree in tall mossy montane 
forest, in association with Phyllocladus, in alpine 
shrubberies, sometimes fire-induced, on margin of 
bog grasslands, 1900-3700 m. Fl. Jan.-Dec, fr. 
July-Jan. Flowers are said to have a slightly fetid 
fragrance. 

Vern. Bolbeh, Chimbu, Masul, gongigl, miluad, 
Chimbu, holai, Asaro, Kefamo, iamuga, Minj, 
Togoba, kumbag, Togoba, kungum, Poio, Enga 
lang., kunguma, Goroka, Togoba, ontkumanip, 
Wahgi, Minj, paiwadedie, Mt Ne, Huli lang., 
paiweriedie, Margarima R., Huli lang., pohn, 
Hagen, Togoba, uinyambangau, Kubor, Minj, 
wanepape, Sirunki, winjabunggawont, Minj, mara, 
ypap, Wabag, Enga lang. 

t. var. revoluta Noot. Leid. Bot. Ser. 1 (1975) 174. 
— Fig. 7. 

Shrub 1-3 m to tree 10 m. Twigs appressedly 
pubescent to villous or tomentose. Leaves 
appressedly sericeous to pubescent or tomentose 
beneath, especially on midrib and nerves, glabres- 
cent, ovate to elliptic, with cuneate to cordate base, 
strongly revolute or recurved margin and rounded 
to acuminate apex, (2V 2 -)4-10 by (l-)2V 2 -6 cm; 
nerves (4-)7-10 pairs; petiole (2— )10— 15 mm. Spike 
basally branched, to 3 cm, becoming much longer 
in fruit, axis densely pubescent to villous or 
tomentose; bracts often broadly boat-shaped, 
3-4 mm. Bracteoles 2 mm, both persistent, 
appressedly long pubescent to villous. Calyx with 
same indument, (nearly) entirely divided into 1-2 
mm long lobes. Corolla 2-4(-5) mm. Stamens 
10-60. Disk glabrous, with few hairs, or densely 
pilose. Ovary with same indument as calyx, 1-2 mm 
high. Style glabrous. Fruit ovoid to ellipsoid, 10-1 1 
by 6-7 mm. Seed more or less curved towards the 



base, embryo from nearly straight to U-shaped. 

D i s t r. - Malesia: New Guinea. 

Ecol. Mossy forest, alpine shrubberies, on 
ridges and in valleys, constituent of subalpine 
forest of Xanthomyrtus, Papuacedrus, Quintinia, 
and Ericaceae, sometimes on peaty soil, 2200- 
3600 m. Fl. Febr.-Aug., fr. July-Dec. Ripe fruit 
purple blue. 

Vern. Bug-bakl, Minj. 

u. var. sogeriensis (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 175. — 5. sogeriensis Brand, Pfl. R. Heft 6 
(1901) 49. — S. angiensis Kaneh. & Hatus. Bot. 
Mag. Tokyo 56 (1942) 485. — Fig. 7. 

Shrub 2-5 m to tree 22 m by 25 cm 0. Twigs 
glabrous (or appressedly pilose in innovations). 
Leaves glabrous, ovate or elliptic, with cuneate to 
rounded base, mostly crenate margin and rounded 
to faintly acuminate apex, 2V2-H by l l /2-7cm; 
nerves 5-9 pairs; petiole 5-20 mm. Spike basally 
branched to c. 3 cm, axis glabrous or appressedly 
pilose. Bracts and bracteoles persistent, glabrous 
or appressedly pilose, V2-IV2 and V2-I mm long 
respectively. Calyx glabrous, or lobes shortly pilose 
towards the apex, V2-IV4 mm, lobes 1 I 2 mm long. 
Corolla 2-3 mm. Stamens less than 10 in ? flowers, 
to 30 in c? flowers. Disk glabrous (or with few hairs). 
Ovary glabrous, i 1 2-^/2 mm high; style glabrous. 
Fruit (ovoid to) ellipsoid, 5-9 by 3-5 mm; stone 
shallowly lengthwise or irregularly grooved. 

Distr. Malesia: New Guinea. 

Ecol. Montane to subalpine rain-forest and 
subalpine scrubberies, in stunted Nothofagus- 
Myrtaceae mossy forest, or forest dominated by 
Castanopsis or by Podocarpus-Papuacedrus, scat- 
tered in subalpine grasslands, on Mt Wilhelmina 
even at 3560 m in sheltered places still a con- 
stituent of 8-10 m high stunted forest; (1950-) 
2100-3560 m. Fl. Sept.-April, fr. Jan.-Nov. Fruit 
turns bluish black when mature. Underside of 
leaves has sometimes globular, pea-sized galls. 

v. var. versteegii (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 176. — S. versteegii Brand, Nova Guinea 
14(1924) 188. 

Shrub or treelet to 5 m. Twigs densely tomentose 
or pilose. Leaves elliptic, except the tomentose or 
pilose midrib and nerves glabrous, or the whole 
surface covered by a cobweb-like or a long-pilose 
indument, base cuneate, apex not or slightly acumi- 
nate to mucronate-caudate, 10-16 by 4-672 cm; 
nerves 6-14 pairs; petiole 6-18 mm. Fascicles in 
the axils of the upper leaves or on wood. Bracts 
and bracteoles persistent, appressedly long pubes- 
cent or sericeous, 4-5 and 2-3 mm long respectively. 
Calyx with same indument, divided into 2-3 mm 
long lobes. Corolla c. 5 mm. Stamens c. 50. Disk 
pilose. Ovary glabrous, 1-2 mm high; style glab- 
rous. Fruit not seen. 

Distr. Malesia: New Guinea. 

Ecol. Rain-forest, 100 and 1300 m. Fl. Febr., 
June-July. 

w. var. maculata (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 176. — 5. maculata Brand in K.Sch. & 
Laut. Nachtr. (1905) 348; Bot. Jahrb. 54 (1916) 
222. — S. margarita Brand, Bot. Jahrb. 54 (1916) 
215. — S. pisifera Brand, I.e. 216, inch var. 
miophylla Brand. — S. ensicuspis Brand, I.e. 219. 



1977] 



Symplocaceae (Nooteboom) 



255 



— 5. arfakensis Gibbs, Arfak (1917) 175. — 
S. morobeensis Sleum. in Fedde, Rep. 42 (1937) 
265. — Fig. 7. 

Shrub 1-2 m to small or moderate tree up to 
15 m by 25 cm 0. Twigs glabrous. Leaves glabrous, 

— elliptic with cuneate, decurrent base, mostly 
entire margin and acuminate apex, 2-13 by l 1 4 -4 
cm ; nerves 4-10 pairs ; petiole 3-15 mm. Spike very 
slender, often branched, 2-10 cm, axis pubescent 
or puberulous to glabrous. Bracts caducous or 
persistent, l-l 1 4 mm long, with the c. 3 / 4 mm long 
bracteoles pubescent or puberulous to glabrous. 
Calyx glabrous, divided into l 4 -l mm long ciliate 
lobes. Corolla 2-4 mm. Stamens from less than 10 
and sterile in ? flowers to 25 in £ flowers. Disk 
pilose. Ovary glabrous, 1 2 -l ' , 4 mm high ; style glab- 
rous. Fruit ovoid to ampulliform, 4-6 by 3-4 mm. 

Distr. Malesia: New Guinea (incl. Sudest, Misi- 
ma & Rossel Is.); common in New Guinea. 

Ecol. Both in the lowland rain-forest at 150- 
300 m (Louisiades) as well as in montane rain-forest 
at 1600-2800 m, where associated with Nothofagus, 
Araucaria and Castanopsis, on narrow crests 
sometimes said to be abundant, also in secondary 
forests. Fl. Aug.-Jan. (June), fr. Aug. -Jan. Ripe 
fruit blue-black. 

Vern. Comogu, Mendi, kunguma, Minj, Togoba, 
mokgeh, Hagen, Togoba lang., ouksanok, Tele- 
fomin. 

x. var. parvifolia Noot. Leid. Bot. Ser. 1 (1975) 
177. — Fig. 7. 

Shrub l 1 2 -4 m to tree up to 10 m, often bushy 
and much-branched. Twigs (appressedly) pubes- 
cent or puberulous. Leaves glabrous, ± elliptic, 
with cuneate, attenuate base, denticulate or dentate 
margin and acute or acuminate apex, l'/4-4 by 
1 . 4 -l J 4 cm; nerves 5-7 pairs; petiole 2-4 mm. Spike 
small, few-flowered, to 1 cm, axis puberulous. 
Bracts and bracteoles persistent, puberulous, 1-2 
and ' 2 -l mm long respectively. Calyx appressedly 
puberulous, divided into c. 1 mm long lobes. 
Corolla 2-2' 2 mm. Stamens c. 10 in y flowers to 25 
in J flowers. Disk densely soft hairy. Ovary glab- 
rous or appressedly puberulous, l-l'/ 2 mm high; 
style glabrous or hairy towards the base. Fruit 
ovoid to ellipsoid, 7-10 by c. 4 mm. 



Distr. Malesia: East New Guinea. 

Ecol. Understorey treelet in lower montane to 
subalpine rain-forest dominated by Nothofagus 
and conifers (Podocarpus and Papuacedrus), often 
mossy, also on forest edges, 1850-3300 m. Fl. 
June-Oct.,/r. Aug. 

Uses. Flush is sometimes eaten as vegetable. 

Vern. Gili, Ebenda, Mendi lang. 

17. Symplocos colombonensis Noot. Leid. Bot. 
Ser. 1 (1975) 177. — Fig. 7, 14a-c. 

Small tree to 10 m. Twigs appressed-pubescent, 
dark brown. Leaves alternate, sparsely appressedly 
pilose beneath, especially on the margin, ovate, 
with cuneate to rounded base often revolute margin 
and acuminate apex, 4-9 by l l l 2 -yi 3 cm; nerves 
7-11 pairs; petiole 3-4 mm. Raceme c. 3-flowered, 
to 3 cm long, axis finely appressedly pubescent. 
Bracts and bracteoles soon caducous, pubescent. 
Pedicel 1-5 mm. Calyx appressedly brown-pilose, 
1 J 4 -3 mm, lobes triangular, l 1 2 -2' 2 mm. Corolla 
glabrous, or thinly red-hairy on the outside in bud, 
c. 5 mm. Stamens e. 90 or more. Disk glabrous or 
with some hairs. Ovary appressedly brown-pilose, 
lV2-2mm high; style glabrous, 4-5 mm. Fruit 
(obliquely) ovoid to ellipsoid, 10-14 by 6mm; 
stone except the apical 2-3 mm brain-like grooved. 
Seed not seen, but embryo probably straight. 

Distr. Malesia: Borneo (Mt Kinabalu). 

Ecol. Mountain forest, 2100-2800 m. Fl. 
Febr.-March, June-July, fr. July, Dec. 

Note. Resembles S. zizyphoides, but a tree with 
less zigzag twigs, larger leaves with longer acumi- 
nate apex, and with calyx lobes longer in propor- 
tion to the tube. 

18. Symplocos composiracemosa Noot. Leid. Bot. 
Ser. 1 (1975) 178. 

Twigs glabrous. Leaves glabrous, elliptic, with 
cuneate, acute base, entire or slightly undulate 
margin and acuminate apex, 8-13'/ 2 by 2'/ 2 -7cm; 
nerves 5-9 pairs, meeting in a looped intramarginal 
vein; petiole 13-15 mm. Raceme compound, to 
5 cm; axis sparsely minutely pilose. Bracts and 
bracteoles persistent, with same indument, I and 
l /a mm long respectively. Pedicels at most 1 mm. 
Calyx glabrous, divided into the rounded, scnu- 





Fig. \4. Symplocos cohmbonensisHooi i Habit, nat. size, b. fruit, c CSof fruit, both ! S.costata 

(hi i (ii'. i., .1 CS of fruit, nat. size. 5. dtflexa Staff, c Habit, nat. size, f. deflorated flower, i 

{a < (iimi-.s J3706, </ Kooaoeas 109* naoou L499X 



256 



Flora Malesiana 



[ser. I, vol. 8 : 



elliptic, recurved, 1 l 1 -l mm long lobes. Corolla 
c. 2 mm. Stamens 15-25, rather stiff. Disk glab- 
rous. Ovary glabrous, c. 1 mm high; style glabrous, 
1 mm. Immature fruit elliptic. 

Distr. Malesia: East New Guinea (Morobe 
Distr.). 

Ecol. Slender substage tree, 1300-1800 m.once 
mentioned in understorey of Nothofagus dominated 
ridge. Fl. Aug., Nov. 

19. Symplocos costata (Bl.) Choisy in Zoll. Syst. 
Verz. 2 (1854) 136; Miq. Fl. Ind. Bat. 1, 2 (1859) 
467; K. & V. Bijdr. 7 (1900) 153; Brand, Pfl. R. 
Heft 6 (1901) 52; Koord. Atlas 2 (1914) t. 380; 
Back. & Bakh. /. Fl. Java 2 (1965) 206; Noot. 
Leid. Bot. Ser. 1 (1975) 179, pi. 8a-d, phot. 1-2. — 
Dicalyx costatus Bl. Bijdr. (1826) 1 1 17. — S. cera- 
sifolia (non Wall, ex DC.) Choisy in Zoll. Syst. 
Verz. 2 (1854) 136; Miq. Fl. Ind. Bat. 1, 2 (1859) 
466, pro stirp. Zoll. — S. caryophylloides Zoll. 
(Syst. Verz. 2, 1854, 136, nomen) Nat. Tijd. N. I. 14 
(1857) 161 ; Miq. Fl. Ind. Bat. 1, 2 (1859) 467. — 
Eugeniodes costatum O. K. Rev. Gen. PI. 2 (1891) 
975. — S. arcuata Brand, Pfl. R. Heft 6 (1901) 58. 
— S. sericea Brand, I.e. 58; Bull. Herb. Boiss. II, 6 
(1906) 748. — Fig. 7, 14d, 15, 16. 

Tree to 20 m, 40 cm 0. Twigs glabrous, often 
with cushion-shaped conspicuous leaf-scars, ter- 
minal buds with many scales, 5-10 mm long. 
Leaves glabrous, narrowly ovate to elliptic, with 
cuneate, acute base, slightly dentate, nearly entire 



margin and acuminate apex, 6-21 by 2-7 cm; 
nerves (8-)10-13(-14) pairs; petiole 10-25 mm. 
Spike from the axils of the leaves or on wood, in 
bud resembling a cone like in S. barringtoniifolia, 
becoming at most 4 cm long, axis tomentose to 
pubescent. Bracts and bracteoles densely sericeous 
to pubescent, broadly boat-shaped, 5-8 mm long, 
soon caducous, and 2-3 mm long, later caducous 
respectively. Calyx glabrous, entirely divided into 
(narrowly) ovate to triangular, 2V 2 -3 mm long 
lobes. Corolla 3-5 mm. Stamens 60 to more than 
100. Disk shortly pilose. Ovary glabrous, c. V2 mm 
high ; style glabrous except sometimes the very base, 
3-6 mm long. Fruit ellipsoid to cylindrical, often 
slightly curved, azure blue, 20-40 by 8-20 mm; 
mesocarp thick, corky, stone with c. 8 high ridges, 
3-celled with a central canal, often only 1 cell 
developed. Seed cylindrical ; embryo straight. 

Distr. Malesia: West & Central Java (E as far 
as G. Telemojo). Fig. 17. 

Ecol. High mountain forest, 900-2000 m, 
scattered. Fl. Aug.-Nov., fr. Aug.-March. 

Vern. Kigledog (Tjibodas), ki telor, ki tomkil, S. 

20. Symplocos crassipes Clarke, Fl. Br. Ind. 3 
(1882) 580; Brand, Pfl. R. Heft 6 (1901) 52; K. & 
G. J. As. Soc. Beng. 74, ii (1906) 245; Ridl. Fl. 
Mai. Pen. 2 (1923) 305; Noot. Leid. Bot. Ser. 1 
(1975) 180, pi. 9-10. 

For synonyms see under the varieties. 

Shrub or small tree to 18 m. Twigs glabrous, or 




Fig. 1 5. Symplocos costata (Bl.) Choisy. Left a tree at Tjibodas Botanic Garden, West Java, 1450 m ; right 
a twig in bud (Nooteboom 885). Photogr. Nooteboom, Febr. 1969. 



1977] 



Symplocaceae (Nooteboom) 



257 




Fig. 16. Symplocos costata (Bl.) Choisy. Close-up 

of flowers in anthesis (Nooteboom 885). Photogr. 

Nooteboom, Febr. 1969. 



(obliquely) pubescent to appressedly or spreadingly 
long-hairy, sometimes with a double indument of a 
short tomentum and long spreading hairs. Leaves 
(narrowly) elliptic to ovate, beneath sparsely 
appressedly pilose, nearly glabrous, to densely 
appressedly to spreadingly long-hairy, rarely also 
hairy above, with cordate to cuneate base, re- 
curved, entire to glandular denticulate margin and 
acuminate apex, 6-27 by 17«-8Va cm; nerves 3-1 1 
pairs; petiole 1-10 mm. Spike short, often clus- 
tered, to l(-2)cm, from the axils of the upper 
leaves, rarely flowers solitary, axis subglabrous to 
appressedly pubescent, or with long, spreading to 
appressed, stiff, brown to rusty hairs 1-4 mm long. 
Bracts and bracteoles persistent, (broadly) ovate, 
triangular or semi-elliptic, rarely acuminate, hairy. 
Calyx hairy or glabrous, whether or not entirely 
divided into the lobes. Corolla 2'/ 2 -6 mm. Stamens 
c. 30 to c. 100. Disk glabrous. Ovary hairy, 1-2 mm 
high; style glabrous, but often with conical, hairy 
base. Fruits mostly 1-2 from each inflorescence, 
glabrous or sparsely long-hairy, bright blue in 
vivo, cylindrical, narrowed towards the apex, 
13-18 by 3-5 mm; stone with c. 12 lengthwise 
grooves; cells 1-3. Seed usually 1, straight with 
straight embryo. 

Distr. Continental Asia (Peninsular Thailand), 
in Malesia: Malay Peninsula (incl. Pcnang) and 
Borneo. 



KI.Y TO THE VARIETIES 

I. Leaves ovate, to 6cm long and 2'/ 4 cm wide; 

nerves 3 6 pairs. Rowen solitary. 

v. vur. havilandii 
I, Leaves (narrowly) ovate to elliptic, 5 1 , 11 by 

2 H'jcm; nerves 3 II pairs. 



2. Leaf-base cordate, base angle 90-180°. 
3. Leaves 5'/2-14 cm long. Petiole 1-2 mm 

b. var. brandiana 

3. Leaves 16-18 cm long. Petiole c. 5 mm. 

a. var. crassipes 
2. Leaf-base not cordate. Base angle 25-90°. 

4. Leaves sparsely appressedly pilose beneath, 
but the indument inconspicuous and leaves 
seemingly glabrous. 

5. Twigs glabrous, rarely appressed-pubescent. 
Calyx often glabrous or nearly so, rarely 
appressed-pubescent. Style-base glabrous, 
rarely pilose c. var. curtisii 

5. Twigs appressed-long-hairy, rarely glabrous. 
Calyx appressed-pubescent. Style-base pilose. 

d. var. ernae 
4. Leaves densely appressed-hairy to sparsely 
more or less appressed-long-hairy beneath, 
indument always evident. 

6. Twigs densely patently brown hairy (hairs 
often c. 2 mm). Leaves sparsely (appressedly) 
long-hairy beneath. Nerves 6-11 pairs. 

f. var. penangiana 

6. Twigs densely obliquely pubescent. Leaves 

densely appressed-pilose beneath. Nerves 

4-6 pairs .... g. var. rufomarginata 

a. var. crassipes. 

Twigs sparsely appressedly long-hairy. Leaves 
sparsely appressed-pilose beneath, the hairs incon- 
spicuous, elliptic, with cordate base and acuminate 
apex, 16-18 by 6-8 cm; nerves c. 10 pairs; petiole 
much swollen, 5 mm. Inflorescence and flowers as 
in var. brandiana (sec. Clarke). 

Distr. Malesia: Malay Peninsula (Johore), only 
known from the type. 

b. var. brandiana (K. & G.) Noot. Leid. Bot. Ser. 1 
(1975) 182. — S. brandiana King & Gamble, J. As. 
Soc. Beng. 74, ii (1906) 245. 

Small tree, 3-8 m. Twigs patently dark brown 
pubescent to tomentose and long-hairy. Leaves 
(appressedly) long-hairy beneath, but midrib and 
nerves patently hairy, with cordate base, 5'/j-14 by 
l 3 / 4 -5 cm; nerves 6-10 pairs; petiole 1-2 mm. Spike 
often on a reduced twig with many cataphylls. 
Bracts and bracteoles narrowly ovate, appressedly 
long-hairy, 3-8 mm. Calyx divided into mate, 
acuminate, appressedly brown hairy, 2'/2-3 mm 
long lobes. Stamens 60 or more. Style with hairy 
conical base, 4 mm. Fruit hairy. 

Distr. Malesia: Malay Peninsula. 

Ecol. Mixed forests, 100 1500 in. 

c. var. curtisii (Oi.iv.) Noot. Leid. Bot. Scr. I (1975) 
183, pi. 9b c. S. curtisii Ouv. In Hook. Ic PL 18 

(1888) t. 1757. — S. montlcola Kim. A: GaMBI i . J. 

, Beng. 74, ii (1906) 235 ;Rtdi . Fl, Mai. Pen. 
2(1923)301. — Fin. 7. 

hcclci "i ihnib to I0m, 35 cm 0. Twigs glab- 
rous or rarely appressed-pubescent. I eaves usually 
sparsely appressedly pilose, neaily glabrous bc- 

neaih. with cuneate, slightly attenuate base. 
H 1 , ih by i B'/icm; nervei A 9 pain; petiole 
3 7 nun. Spike contracted, often branched, axil 
glabrous to eppressedl) pubescent. Bracts and 
bracteoles ovate to triangular, appressedly pubes 

Cent, 11'; and < 1 nun long respectively. < .dvx 



258 



Flora Malesiana 



[ser. I, vol. 8 2 



glabrous or nearly so, rarely appressedly pubes- 
cent, lV2-2mm, the lobes 1 l 2 -\ l l 2 Tnm, becoming 
longer by tearing apart when older. Corolla 
372-4 mm. Disk glabrous. Ovary appressedly 
pubescent, often narrowly funnel-shaped, l l l 2 -2 
mm high; style glabrous, the base glabrous or 
pilose. Fruit glabrous, deep blue. 

Distr. Continental Asia (Peninsular Thailand), 
in Malesia: Malay Peninsula (Johore, Selangor). 

Ecol. Hill rain-forest, 200-1400 m. Fl. Aug.- 
Jan.,/r. Febr.-May, Oct., flowers scented. 

Vern. Malaya: kayu jenerku, Selangor: Temuan. 

d. var. ernae (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 184, pi. 10b. — S. ernae Brand, Pfl. R. 
Heft 6 (1901) 58; Merr. En. Born. (1921) 486. — 
Fig. 7. 

Shrub or slender tree to 18 m, 15 cm 0. Twigs 
appressedly (long-)hairy, rarely glabrous. Leaves 
sparsely appressedly pilose, nearly glabrous 
beneath, with cuneate base, 6-15(-18) by 2 1 / 2 -6 
(-7) cm; nerves 3-6 pairs; petiole 3-5 mm. Spike 
basally branched, contracted, axis appressedly 
pubescent. Bracts and bracteoles broadly ovate, 
often boat-shaped, appressedly pubescent, c. 1 mm 
long. Calyx appressedly pubescent, 1 V 4 -2 mm long, 
lobes l-l'/ 2 mm, often becoming longer in older 
stage. Corolla 3-5 mm. Stamens c. 30 to c. 70. 
Disk glabrous. Ovary appressedly pubescent, 
l-lV2mm high; style glabrous. Fruit glabrous. 

Distr. Malesia: Borneo (Sarawak, Brunei, 
Sabah; also in W. Kutei: G. Kemul). 

Ecol. Lowland mixed Dipterocarp forest, also 
in a swamp forest, and in hill rain-forest on sandy 
clay, from sea-level to 1500 m. Fl. Sept.-Oct., 
Febr.-June, fr. July, Nov. 

e. var. havilandii (Brand) Noot. Leid. Bot. Ser. 1 
(1975) 184, pi. 10c. — S. havilandii Brand, Pfl. 
R. Heft 6 (1901) 41 ; Merr. En. Born. (1921) 486. 

Treelet. Twigs pubescent. Leaves ovate, rather 
densely appressed-pilose, especially on midrib and 
nerves and along the margin, with rounded base, 
2 3 / 4 -6 by 1 V4-2 3 / 4 cm ; nerves 3-6 pairs ; petiole 2-3 
mm. Flowers solitary, sessile from the axils of the 
leaves. Bracts and bracteoles appressedly (long-) 
pubescent, semi-orbicular 2 mm long and ovate 
I 1 1 2 mm long respectively. Calyx appressedly 
pubescent, 2 mm, the lobes P/2 mm long. Corolla 
Z*/a mm. Stamens c. 35. Disk glabrous. Ovary 
appressedly pubescent, c. 1 mm high; style with 
pilose base. Fruit pale blue. 

Distr. Malesia: Borneo (Sarawak). 

Ecol. Hill rain-forest, 600-900 m. Fl.fr. July. 

f. var. penangiana (K. & G.) Noot. Leid. Bot. Ser. 1 
(1975) 185, pi. 9d. — S. penangiana King & 
Gamble, J. As. Soc. Beng. 74, ii (1906) 245; Ridl. 
Fl. Mai. Pen. 2 (1923) 306. — Fig. 7. 

Shrub or treelet to 10 m. Twigs densely patently 
dark brown hairy. Leaves narrowly elliptic, sparsely 
(appressedly) long-hairy beneath, especially on 
midrib and nerves, rarely also long-hairy above, 
with rounded to acute base, 6-27 by 2V 2 -8 cm, 
margin often sharply glandular dentate, appres- 
sedly long-hairy beneath; nerves 6-11 pairs; 
petiole 2-10 mm. Spike contracted, branched, axis 
densely more or less appressedly villous, hairs 
1-4 mm. Bracts and bracteoles appressedly dark 



brown long-hairy, narrowly elliptic to ovate, 
sometimes caudate, 1-7 mm. Calyx densely 
appressedly dark brown hairy, entirely divided into 
1 V2-4 mm long lobes. Corolla 3-6 mm. Stamens 
30 to more than 100. Disk pilose. Ovary with same 
indument as calyx, l-l'/^mm high; style with 
pilose base. Fruits hairy, pink. 

Distr. Malesia: Malay Peninsula (incl. Penang). 

Ecol. Lowland rain-forest, 150-500 m. Fl. May, 
fr. Nov., April. 

g. var. rufomarginata Noot. Leid. Bot. Ser. 1 (1975) 
185, pi. 10a. 

Shrub or treelet to 5 m. Twigs densely pubescent. 
Leaves rather densely appressedly hairy beneath, 
ovate to elliptic, with cuneate base and margin 
densely appressedly rufous-hairy beneath, 5*/r- 
IIV2 by 2-3V2cm; nerves 4-6 pairs; petiole 
2-3 mm. Spike much contracted, axis hairy. 
Bracts and bracteoles (broadly) elliptic, c. 3 mm. 
Calyx densely, appressedly long sericeously pubes- 
cent, entirely divided into 2 mm long rounded 
lobes. Corolla c. 2 l / 2 mm. Stamens c. 25. Disk 
glabrous. Ovary with same indument as calyx, c. 

1 mm high; style hairy halfway up. 

Distr. Malesia: Borneo (Sarawak, near Kuch- 
ing). 

21. Symplocos cylindracea Noot. Leid. Bot. Ser. 1 
(1975) 187. — Fig. 7. 

Tree 10-30 m, 35 cm 0. Twigs glabrous, or 
pubescent in innovation. Leaves glabrous, or mid- 
rib (and nerves) minutely hairy beneath, ± elliptic, 
acuminate with acute to rounded, attenuate base 
and crenate or crenulate margin, 9-15 by 3V2- 
9V2 cm ; nerves 6-9 pairs, meeting in a looped 
intramarginal vein ; petiole 7-20 mm. Flowers in an 
up to 8 cm long panicle with minute or shortly 
pilose axis. Bracts and bracteoles caducous, glab- 
rous or minutely hairy, ciliate, ovate, c. 3 and c. 

2 mm long respectively. Pedicel l /r~3 mm, some- 
times seemingly much longer when only one flower 
is left on a small branch. Calyx 2-V-\ 2 mm long, 
entirely divided into elliptic to nearly semi-orbi- 
cular lobes, sparsely appressedly pilose to glabrous, 
ciliate. Corolla 5-6 mm. Stamens more than 100. 
Disk 5-glandular, pilose except the glands. Ovary 
glabrous or pubescent, l-l'/ 2 mm high; style 
(minutely) pilose, 2-4 mm. Fruit cylindrical, 15 by 
5-6 mm, mesocarp fleshy, stone with low length- 
wise ridges, 3-celled, 1, 2, or all 3 cells developed. 
Seed 1 in each fertile cell, straight with straight 
embryo. 

Distr. Malesia: New Guinea (West and North, 
Morobe Distr., Central Div., and New Britain). 

Ecol. Plain rain-forest, also in Anisoptera forest 
on ridge top, 60-800 m. Fl. Jan.-July, fr. Febr.- 
March. 

22. Symplocos deflexa Stapf, Trans. Linn. Soc. 4 
(1894) 205; Brand, Pfl. R. Heft 6 (1901) 64; 
Gibbs, J. Linn. Soc. Bot. 42 (1914) 109; Merr. En. 
Born. (1921) 487; Noot. Leid. Bot. Ser. 1 (1975) 
188. — Fig. 7, 14e-f. 

Treelet to 6 m high and 8 cm 0. Twigs obliquely- 
patently brown hairy, ± zigzag. Leaves alternate, 
glabrous above, rather densely pilose beneath, 
especially towards the margin, elliptic, acuminate, 
obtuse or acute, with rounded or sharply attenuate 



1977] 



Symplocaceae (Nooteboom) 



259 



base and recurved to revolute, sharply glandular 
dentate margin, 3-5 by l 1 2 -2' 2 cm; nerves 5-7 
pairs, usually merging into the reticulation; petiole 
1-2 mm, densely patently brown hairy. Flowers 
fragrant, in an up to 6-flowered, 1-4 cm long lax 
raceme which is appressedly to patently brown 
pilose in all parts except the corolla. Bracts and 
bracteoles persistent, c . 5 by 3 and c. 3 by l 1 2 mm 
respectively. Pedicel 2-4 mm. Calyx divided into 
obtuse and semi-elliptic to acute and triangular 
lobes, c . 1 ' 2 mm long. Petals 5-7, glabrous or the 
outer ones minutely appressedly hairy, 4-6 mm 
long. Stamens 60-90. Disk low, 5-glandular, 
sparsely long-pilose. Ovary lV 2 -2mm high; style 
c. 5 mm, gradually thickened towards its base, the 
lower half sparsely long-pilose. Fruit ovoid, often 
curved, including the persistent calyx c. 10 by 5 mm ; 
stone c. 8 by 4 mm with shallow grooves and large 
apical pore. Seed straight with straight embryo. 

Distr. Malesia: Borneo (Sabah, only found on 
Mt Kinabalu near Paka cave). 

Ecol. Low subalpine forest and mountain 
scrub, 2400-3200 m. Fl. Oct.-Febr., fr. March, 
Aug.-Oct. 

23. Symplocos fasciculata Zoll. Syst. Verz. 2 
(1854) 136; Nat. Tijd. N. I. 14 (1857) 161; Miq. 
Fl. Ind. Bat. 1, 2 (1859) 467; Suppl. 1 (1861) 474, 
incl. var. minor Miq. I.e. 475 ; Clarke, Fl. Br. Ind. 3 
(1882) 574; K. & V. Bijdr. 7 (1900) 150, incl. var. 
blumeana K. & V. I.e. 151 ; Brand, Pfl. R. Heft 6 
(1901) 34; K. & G. J. As. Soc. Beng. 74, ii (1906) 
235; Koord. Atlas 2 (1914) t. 383; Ridl. Fl. Mai. 
Pen. 2 (1923) 301; Heyne, Nutt. PI. (1927) 1262; 
Merr. Un. Cal. Publ. Bot. 15 (1929) 248; Burk. 
Diet. (1935) 2113; Corner, Ways. Trees (1940) 622, 
t. 231; Back. & Bakh. /. Fl. Java 2 (1965) 205; 
Noot. Leid. Bot. Ser. 1 (1975) 191, f. 2c, pi. 13. — 
Sariava Reinw. Syll. Ratisb. 2 (1825) 12. — 
Dicalyx tinctorius Bl. Bijdr. (1826) 1116, non 
S. tinctoria L'Herit. 1791. — Eugeniodes fascicu- 
latum O. K. Rev. Gen. PI. 2 (1891) 409. — S. 
phanerophlebia Merr. Philip. J. Sc. 9 (1914) Bot. 
382; J. Str. Br. R. As. Soc. n. 76 (1917) 112; En. 
Philip. 3(1923)301. — Fig. 7. 

Shrub, or less often a tree to 22 m high and 50 
cm 0. Twigs sparsely pilose, puberulous, or appres- 
sedly pubescent, glabrescent, often zigzag. Leaves 
alternately or (on the leaders) spirally arranged, 
glabrous above, sparsely appressedly fine-hairy 
beneath, rarely patently hirsute, especially on mid- 
rib and nerves and towards the margin, (narrowly) 
elliptic or sometimes ovate, acuminate to caudate 
with acute to rounded base, 5—1 3C-1 8> by 2 4' , 
(-6) cm; nerves (4-)6 H( 1 1 ) pairs, meeting in a 
looped intramarginal vein ; petiole 2 8 mm. Flowers 
in a fascicle of reduced, often branched, racemes to 
2 1 j cm long. Bracts and bracteoles persistent, 
minute (rarely to 3mm), as the axis pubescent; 
often several bracts present, indicating the origin 
from a more branched inflorescence, Pedicel 
1 5 mm, pubescent. Calyx divided into (4 )5( 6) 
broadly ovoid, rounded, apprcsscdk pubescent or 

f;labrous lobes, r. 1 mm long but sometimes the- 
ories different in si/e. often some of the lobes 
petaloid. < orolltt glabrous or more olten with 
minute hairs towards the outer base, raiely some 
n the back too, 2 4 1 , mm Stamens 12 35. 
Disk glabrous to more or less pilose, low annular. 



Ovary appressedly hairy, c. 1 mm high ; style hairy, 
especially towards the thickened base, rarely 
glabrous, 2-3 1 : 2 mm. Fruit broadly or narrowly 
ampulliform, often curved, the belly globose or 
ovoid, the neck broadly conical, dark violet-blue 
or cobalt-blue, 5-7 by 3-5 mm; stone brain-like 
grooved without or with c. 10 shallow grooves. 
Seed 1, much lobed, with slightly curved embryo. 
Distr. Extreme South Peninsular Thailand 
(Pattani) and throughout Malesia, except the 
Lesser Sunda Is., the Moluccas, and New Guinea. 
One of the most common Symplocos species in 
Malesia. Fig. 17. 




Fig. 17. Ranges of a. Symplocos costata (Bl.) 

Choisy, b. S. cerasifolia Wall, ex DC. var. cerasi- 

folia, c. S. fasciculata Zoll. 



Ecol. In primary high and open secondary 
forest and thickets, common in disturbed forest, 
rather indifferent to soils, besides on latosols, 
recorded from sand (Banka), in Borneo from sand- 
stone, black soils, seasonally swampy land and 
Dipterocarp forest, also riparian, in Udjong Kulon 
from raised coral limestone, from sea-level up to 
c. 2200 m. Fl. June-Sept. (Nov.-April), fr. Sept.- 
March. Several times flowers are noted to be scent- 
ed, but once recorded as emitting a pervasive sour 
smell (Malaya, Whitmore). 

Vern. Malaya : kCrCnang, nasi-nasi, miruui 
(obviously referring to the often unripe white 
fruit, resembling grains of cooked rice), M, Kepong, 
ttbiak, Sclangor; Sumatra: kavu lohu-lohu, 
Asahan, djauik bulau, I'ajakumhu. djunk, Kerintji. 
ktkatja, lilibah, Bengfcalis, pipi udiin, kaio. 
Irhomclukut, M. Ulu. I'alembang, luipii-liiipii, 

havu-havu, h. h. dibk, //. h. itam, ft. h. udtng, 
kareui karnd udbig, lihat-lihai uding, Simalur, 
I-;,;. Kepahiang, kayu lebeu. PaJembang, djarok, 

Hank, i ; Java; dpimk, ,////« A, ,/. Irnlik. </ pi it, 

d uiuii'.d wulu,J,S,klptit,8; Borneo s.iiawak: 
flrah. [ban, pirlaboh, Murut; Sabah: labah, Uboh, 
lobon, KJnabatanaan, Kadaaan lang., gtak, 
in, Idaho, Duiun; Brunei: pachal ambok; 
Kalimantan njam-njam, Buhu&n,fwtdti/tiputih, 
Balikpapan. 



260 



Flora Malesiana 



[ser. I, vol. 8 2 



Notes. The fruit is of a type usually containing a 
curved seed with curved embryo; here it is, how- 
ever, only slightly curved. 

In the herbarium sterile sheets are sometimes 
confused with Eurya acuminata which has, in 
Malaya, often the same vernacular names; cf. 
Corner (1940). 

Normally lateral shoots are collected which have 
a characteristic alternate phyllotaxis, but I have 
also found leader-shoots which have a spiral phyl- 
lotaxis flowering in Borneo. 

In habit S. fascieulata is very similar to S. 
laeteviridis but its flowers are truly fascicled with 
more than 3 bracts under each flower and these 
persistent, a regular 5-lobed calyx, an ampulliform 
fruit with a ruminate seed and curved embryo. 
In S. laeteviridis the inflorescence is a raceme or 
panicle with 1 bract and 2 bracteoles under each 
flower and these caducous, a calyx which splits 
into a 3-lobed and a 2-lobed part, while the fruit is 
ellipsoid to ovate, with a non-ruminate seed and a 
straight embryo. 

24. Symplocos filipes Noot. Leid. Bot. Ser. 1 
(1975) 193, pi. 14a-d. — Fig. 7. 

Twigs glabrous or sparsely pulverulent-puberu- 
lous, the terminal buds small, with pulverulent- 
puberulous scales which often bear large vesicular 
glands on the margin. Leaves glabrous or sparsely 
pulverulent-puberulous beneath, ± elliptic, long 
acuminate, with acute often attenuate base and 
entire or slightly denticulate margin which contains 
a row of large vesicular glands, 4 l l 2 -l l j 2 by 2-3 cm ; 
nerves 5-6 pairs, meeting in a looped intramarginal 
vein; petiole 7-8 mm. Flowers in a lax raceme of 
4-10 cm, the axis sparsely pulverulent-puberulous. 
Bracts and bracteoles persistent, with same 
indument, x \ 2 and 1 mm long respectively. Pedicel 
slender, 2—15 mm. Calyx sparsely pulverulent- 
puberulous, divided into semi-ellliptic l [ 2 mm long 
lobes. Corolla c. 3 mm. Stamens c. 25. Disk annu- 
lar, glabrous. Ovary with same indument as calyx, 
c. P^mm high; style glabrous, c. 3 mm. Fruit 
ellipsoid, c. 10 by 4 mm, the small calyx incurved; 
stone spindle-shaped, with shallow lengthwise 
grooves, 1-celled. Seed 1, straight with straight 
embryo. 

Distr. Malesia: Philippines (Mindoro: Mt 
Halcon), two collections. 

25. Symplocos gambliana Brand, Bull. Herb. 
Boiss. II, 6 (1906) 748; Merr. En. Born. (1921) 
484; Noot. Leid. Bot. Ser. 1 (1975) 195. — S. havi- 
landii King & Gamble, J. As. Soc. Beng. 74, ii 
(1906) 251, non Brand, 1901. 

Twigs glabrous. Leaves glabrous, ± elliptic, 
abruptly oblique acuminate with acute, attenuate 
base and entire, recurved margin, 6-9 by 3-4V2 cm ; 
nerves 6-8 pairs meeting in a looped, faintly 
prominent intramarginal vein; petiole 5-10 mm. 
Flowers in a lax spike or raceme to 6 cm; axis 
glabrous. Bracts and bracteoles ?minute, soon 
caducous. Pedicel less than 1 mm. Calyx entirely 
divided into semi-orbicular, ciliate, 3 U-\ 1 l 2 mm 
long lobes. Corolla ciliolate, often with some minute 
hairs on the outside, c. 5 mm. Stamens c. 50. Disk 
5-glandular, with the style base minutely pilose. 
Ovary glabrous, c. 1 mm high; style glabrous 
except the base, 4 mm. Fruit not known. 



Distr. Malesia: Borneo (Sarawak), only known 
from the type. 

26. Symplocos gigantifolia Noot. Leid. Bot. Ser. 1 
(1975) 195. 

Twigs glabrous, very thick. Leaves glabrous, 
obovate, shortly acuminate, the base cuneate but 
truncate at its lowermost part, margin ± entire, 
21-62 by 7-19 cm; nerves 13-20 pairs, merging 
into the venation; petiole c. 1 cm. Flowers in a 
fascicle or very short spike on wood. Bracts and 
bracteoles persistent, appressedly pubescent, semi- 
elliptic, rounded, 1-2 mm. Calyx minutely ap- 
pressedly pubescent, 2 mm, the 3 semi-elliptic, 
rounded lobes c. 1 l j 2 mm long. Corolla 4-5 mm. 
Stamens c. 50. Disk 5-glandular, glabrous, but 
style base pilose. Ovary with same indument as 
calyx, c. 1 mm high; style glabrous, reduced (only 
3 flowers seen). Fruit very young. Seeds not seen. 

Distr. Malesia: East New Guinea (Central 
Division, Southern Highlands and Western Dis- 
trict), 3 collections. 

Ecol. In high forest, once along a riverbed, 90, 
500, and 800 m. Fl. April-May. 

Notes. Brass (3894) noted that it is a 'striking 
tree with erect branching habit and flowers between 
the whorls.' In the three collections studied the 
'whorled' position of the leaves could not be 
checked. Possibly the main leaves may be conspi- 
cuously crowded at the end of the year's growth 
(flush). 

A similar situation is reported to occur in S. 
herzogii, which is the closest related species, 
differing in having smaller, hairy leaves, hairy 
twigs, and larger bracts. 

27. Symplocos glabriramifera Noot. Leid. Bot. 
Ser. 1 (1975) 196, pi. 15a-d. — Fig. 7. 

Twigs glabrous. Leaves glabrous, elliptic to 
obovate, (faintly) acuminate, with acute, attenuate 
base and crenate or crenulate apex, 4-672 by l 1 /*- 
2V2 cm ; nerves 6-8 pairs, meeting in a looped 
intramarginal vein; petiole 5-7 mm. Flowers in a 
short lax raceme to 1 1 I 2 cm, axis glabrous. Bracts 
and bracteoles caducous, glabrous, ciliolate, P/2 
and 1 mm long respectively. Pedicel 1-2 mm. Calyx 
glabrous, c. 1V 2 mm long, the lobes 3, semi-elliptic, 
rounded, c. lV-tmm long. Corolla probably 3- 
merous, 3-4 mm. Stamens 30-50. Disk glabrous, 
3-5-glandular. Ovary glabrous, c. 1 mm high; 
style glabrous. Fruit ellipsoid, truncate at both 
ends, 8-12 by 4-6 mm; stone shallowly lengthwise 
grooved without, 3-celled. Seed 1 in each cell, 
straight with straight embryo. 

Distr. Malesia: Philippines (Luzon: Benguet & 
Nueva Vizcaya Prov.). 

Ecol. Mountain forest, 1900 m. Fl. Febr., May. 

28. Symplocos glomerata King ex Clarke, Fl. Br. 
Ind. 3 (1882) 577; Brand, Pfl. R. Heft 6 (1901) 69; 
Brandis, Ind. Trees (1906) 438; Hand.-Mazz. 
Beih. Bot. Centralbl. 62 B (1943) 30; Noot. Leid. 
Bot. Ser. 1 (1975) 199, pi. 16a-b, with full syno- 
nymy. — Fig. 7. 

var. glomerata. 

Small tree, 6 m. Twigs glabrous, or tomentellous 
and then soon glabrescent. Leaves elliptic, acu- 
minate, with glandular dentate margin, 7-20 by 



1977] 



Symplocaceae (Nooteboom) 



261 



2— 4 1 2 cm; nerves 10-16 pairs meeting in a looped 
intramarginal vein; petiole 5-12 mm. Flowers in a 
fascicle from the axils of the leaves or from wood. 
Calyx glabrous, 1-2 mm, the ciliate lobes slightly 
shorter. Corolla 4-5 mm. Stamens c. 25 to c. 50. 
Disk cylindrical, c. 1 mm high. Ovary glabrous, 
c. 1 mm high. Fruit 7-10 by c. 3 mm. 

Distr. Continental Asia (India, Burma, Indo- 
China, China, Hainan, Hong Kong, Formosa); in 
Malesia: Malay Peninsula (Trengganu, once found 
on G. Lawut Besut). 

Ecol. Montane forest, 1500 m. Fr. April. 

Note. There is a considerable synonymy in- 
volved in this widely spread continental SE. Asian 
species which I have subdivided into two subspecies 
and several varieties. 

29. Symplocos goodeniacea Noot. Leid. Bot. Ser. 1 
(1975)204. 

Small tree to V/ 2 m. Twigs glabrous. Leaves 
narrowly elliptic, shortly acuminate with cuneate 
base and recurved entire or denticulate margin, 
17-30 by 3 l / 2 -7 cm; nerves 11-13 pairs, at least in 
the apical part of the leaf meeting in a looped 
intramarginal vein close to the margin; petiole 
15-25 mm. Flowers in a spike to 4 cm ; axis puberu- 
lous. Bracts and bracteoles persistent, glabrous but 
ciliate, c. 2 mm. Calyx glabrous, divided into the 
broadly rounded l' 2 -2mm long lobes. Corolla 
6-8 mm. Stamens more than 100. Disk annular, 
minutely pilose. Ovary glabrous, l'/ 2 -2mm high; 
style glabrous. Fruit not known. 

Distr. Malesia: Borneo (Sabah), only known 
from the type. 

Ecol. Lowland rain-forest, 150 m. 

30. Symplocos herzogii Sleum. in Fedde, Rep. 42 
(1937) 264; Noot. Leid. Bot. Ser. 1 (1975) 207. — 
Fig. 7. 

Small tree or leaning shrub, 4-6 m high. Twigs 
thick, densely tomentose. Leaves pseudoverticillate, 
but between the whorls the scars of fallen spirally 
arranged leaves visible in at least one collection, 
rather densely hairy beneath, especially on midrib 
and nerves, ± elliptic, acute to acuminate with 
cuneate base (the very base truncate) and sharply 
dentate margin, 13-20 by 5-9'/ 2 cm; nerves 10-17 
pairs; petiole with same indument as twigs, very 
thick, 7-20 mm. Flowers in a fascicle or spike to 
2 cm from the axils of the leaves or from wood. 
Bracts and bracteoles persistent, densely redbrown 
sericeous, c. 5 mm and c. 3 mm respectively. 
Calyx appressedly redbrown hairy, 2-2" 2 mm, the 
lobes : ovate, acute, l', 2 -2 mm. Corolla 3-4 mm. 
Stamens c. 40 in $ flowers (according to Sleumer 
I.e. absent in dossers). Disk pilose. Ovary 
glabrous, ' 2 I mm high; style reduced in ; flowers, 
in . flowers 3' 2 mm (according to SLEUMER I.e.). 
Fruit globose to ampulliform, c. 8 by 6 mm, the 
stone ribbed. Seed I, curved with curved embryo. 

Distr. Malesia last New Guinea (Morobc 
■■■)■ 

I. Midmountain rain-forest, 1500 IKOOm. 
//. Dec. April. 

.ive only seen ; llosseis and fruits. 

dingtoSLEi Mm Boirerearefew,atthebasc 
<>f i he inflorescence. 

This species is allied to S. eig<mt\folUX\ see the 
notes under that species. 



31. Symplocos johniana Stapf, Trans. Linn. Soc. 
Bot. 4 (1894) 206; Brand, Pfl. R. Heft 6 (1901) 65; 
Merr. En. Born. (1921) 487; H. Heine, Pfl. 
Samml. Clemens Kinabalu (1953) 88; Noot. Leid. 
Bot. Ser. 1 (1975) 208, pi. 17f-g. — Fig. 7. 

Shrub or small tree, to 3 m. Twigs densely 
obliquely to patently rusty hirsute. Leaves spirally 
arranged or alternate, rather densely patently 
hirsute beneath, or only midrib and nerves hairy, 
acuminate to caudate with rounded to cordate base 
and usually rather coarsely sharp-dentate margin, 
ovate, 2 1 2 -7 by l 1 / 4 -3 1 / 2 cm; nerves 3-6 pairs 
meeting in a looped intramarginal vein; petiole 
1-2 mm. Flowers in 1-flowered raceme, axis V 2 mm, 
with l'/ 2 mm long bract and the c. 1 mm long 
bracteoles loosely appressedly rusty hirsute. 
Pedicel c. 1 mm. Calyx rusty hirsute, divided into 
the semi-elliptic rounded 1-1 \ 2 mm long lobes. 
Corolla c. 5 mm. Stamens 60-90. Disk stellate, 
densely hirsute. Ovary rusty hirsute, c. 1 mm high; 
style glabrous, c. 6 mm. Fruit narrowly flask- 
shaped, often sparsely hairy, intense indigo-blue, 
c. 13 by 4 mm, the persistent calyx not included. 
Seed 1, straight, narrowly elliptic, embryo straight. 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu; 
W. Kutei: G. Kemul). 

Ecol. In forest, in damp shady places, often in 
crevices of granite rocks, 1500-3200 m. Fl. Febr.- 
Oct.,/r. Febr.-May, Sept.-Oct. 

Note. The distribution is interesting because this 
species, which was assumed to be a Kinabalu 
endemic, is also found on an old, worn-down, 
rather low summit in W. Kutei, in a mountain 
range which is probably older than Mt Kinabalu. 
This feature is also found in some other mountain 
plants, e.g. Lobelia borneensis, which were found 
on Mt Murud, in Sarawak (cf. Steen. Proc. R. Soc. 
Lond. B 161, 1964, 16). Van Steenis concluded 
that Kinabalu plants possibly in the past had a 
wider distribution in Borneo when there were more 
higher peaks in the island, and that the few present 
stations on the low mountains are relict stations 
(cf. also Steen. Mai. Nat. J. 20, 1967, 39). 

32. Symplocos junghuhnii Koord. Proc. Kon. 
Acad. Wet. A'dam 10(1908) 160; Noot. Leid. Bot. 
Ser. 1 (1975) 209. — Fig. 7, 18. 

Twigs glabrous. Leaves glabrous, or with some 
appressed hairs beneath, acuminate with cuneate to 
cordate base and entire to denticulate margin, 
obovate to elliptic, 9 13 by 4'/ 2 -5cm; nerves 
7-10 pairs; petiole 10 17 mm. I lowers in a raceme 
to 6 cm, axis pubescent to tomentose, glabrescent. 
Bracts and bracteoles soon caducous, not seen. 
Calyx glabrous, divided into semi-orbicular 

COrdatdy based < . 2 mm long lobes. Corolla S 10 
mm. Stamens more than KM). Disk 5-glandular, 
with the bio. nils conical style base soft hairy. 
Ovary tomentose, 2 i mm high; stsle glabrous, 

C. 7 mm. hint (only soung fruits seen) : elliptic, 

is hv 8 nun I mbryo probably straight. 

Dim i. Malesia West Jas.i ( I'lcam'ci I jigCD 

teng). 
i "i Mixed montane rain-forest, 1750m. 

Note It is not deal whs this species ssas omitted 

from Ba< k .^ Bakm / .' i I lore ol Is 

13, Svmploius lacUsiridis STAFF, lians. linn SOC 

Bot. 4(1894)205; Bhand, Pfl. R. Heft 6 (1901) 53; 



262 



Flora Malesiana 



[ser. I, vol. 8 2 




Fig. 18. Symplocos junghuhnii Koord. a. Leaf 

underside, nat. size, b. deflorated flower from 

above, showing 5-glandular disk, x 4, c. fruit, x 2 

(Koorders 26420). 

Merr. En. Born. (1921) 487; Airy Shaw, Kew 
Bull. (1939) 408; H. Heine, Pfl. Samml. Clemens 
Kinabalu (1953) 88; Noot. Leid. Bot. Ser. 1 (1975) 
208, pi. 18-19. — Fig. 3. 

For synonyms see under the varieties. 

Shrub or tree to 10(— 21) m. Twigs glabrous or 
clothed by a much variable indument, often faintly 
zigzag. Leaves alternate, glabrous to more or less 
pilose beneath, acuminate to caudate with acute to 



cordate base and nearly entire finely glandular 
dentate or sharply dentate, flat or recurved, margin, 
(narrowly) ovate to elliptic, l 3 / 4 -12 by l-4V 2 cm; 
nerves (3— )4— 1 1 pairs, usually meeting in a looped 
intramarginal vein. Flowers in a raceme or panicle 
to 4 l / 2 cm, the axis clothed with hairs. Bracts and 
bracteoles hairy, soon caducous. Pedicels 0-5 mm. 
Calyx glabrous or hairy, 2-3 mm long, symmetri- 
cally cleft, the lobes 1-3 mm. Corolla 3-5 mm, 
often with minute hairs on the outside. Stamens 
25-70. Disk 5-stellate, shortly minutely pilose. 
Ovary (appressedly) hairy, l-F^rnm high; style 
glabrous, as long as the corolla. Fruit white to 
bluish-black, (obliquely) ovoid to ellipsoid, 7-12 
by (3-)5-6 mm. Seed 1, cylindrical to ellipsoidal or 
ovoid with straight embryo. 

Distr. Malesia: N. Sumatra, Malaya, Borneo, 
and Celebes. 

Note. See for differences with S. fasciculata 
under that species. 



key to the varieties 

1 . Leaf base distinctly cordate. 
2. Twigs with an indument of c. 2 mm long hairs. 
Leaves 5-12 cm long . . . e. var. mjobergii 

2. Twigs with an indument of V4-I mm long hairs. 
Leaves l 3 / 4 -4V2cm long. d. var. kinabaluensis 

1. Leaf base cuneate to rounded. 

3. Twigs velutinous. 

4. Leaves c. 4 cm long . . . f. var. pauciHora 

4. Leaves 9-12 cm long. . . g. var. velutinosa 
3. Twigs glabrous, or pubescent, hairs much 

shorter than 2 mm. 

5. Twigs glabrous or appressed-pubescent. 
Nerves 6-9 pairs. ... a. var. laeteviridis 

5. Twigs loosely appressed-pubescent. Nerves 

3-6 pairs b. var. alternifolia 

3. Twigs obliquely to patently long-pilose, hairs 
of the indument c. 2 mm long. 

c. var. basirotunda 

a. var. laeteviridis. — Cf. Noot. Leid. Bot. Ser. 1 
(1975) 211, pi. 18e-f, 19b. — S. forbesii Brand, 
Pfl. R. Heft 6 (1901) 63. — Fig. 3, 7. 

Shrub or tree to 10(-21) m. Twigs glabrous or 
appressedly pubescent. Leaves often yellowish 
green above, brownish beneath in sicco, acuminate 
to caudate with cuneate to rounded base (narrowly) 
elliptic to ovate, 4-11 by lV2-4cm; nerves 6-9 
pairs, usually meeting in a looped intramarginal 
vein; petiole 1— 3(— 4) mm. Flowers in a predomi- 
nantly basally branched, often very short lax 
panicle of racemes, rarely a simple raceme, to 3 cm 
long; axis pubescent. Bracts and the 0-1 bracteoles 
very soon caducous. Pedicel with same indument as 
axis, 0-2(-5) mm. Fruit black-blue. 

Distr. Malesia: N. Sumatra, Banka, Malay 
Peninsula (Perak, once), Borneo (throughout, 
many collections from Mt Kinabalu), SW. 
Celebes (Bonthain, Todjambu). 

Ecol. In hill and montane rain-forest, in a 
variable set of conditions, on rich clay in mixed 
Dipterocarp forest near a river, on stony hillsides, 
on black soil on ridge top, on a basalt ridge under 
Dipterocarp forest (Sarawak), and even on ultra- 
basic; 500-2000 m. Fl. Jan.-Oct., fr. almost Jan.- 
Dec. 



1977] 



Symplocaceae (Nooteboom) 



263 



Vern. Sumatra: alleban, Karolands, kayu loba- 
loba, k. sae-sae, Asahan; Borneo: Sarawak: luroh, 
Kayan. 

b. var. alternifolia Noot. Leid. Bot. Ser. 1 (1975) 

211, pi. 18a. 

Shrub or treelet. Twigs densely loosely appres- 
sedly brown-pubescent. Leaves rather densely to 
sparsely appressed-pilose beneath, especially on the 
margin, acuminate to caudate with cuneate shortly 
attenuate base and ciliate, recurved, entire to 
finely glandular dentate margin, ± elliptic, 4-5 \ 2 
by l 1 2 -2' 2 cm; nerves (3-)4-6 pairs, meeting in a 
looped intramarginal vein but sometimes obscured 
by the indument; petiole c. 2 mm. Flowers in a 
(sometimes branched) raceme to 3 cm or solitary, 
axis red-brown pilose. Pedicel 0-' 2 mm (to 4 mm 
in solitary flowers). 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu). 

Ecol. Montane rain-forest, 1000-1500 m. Fl. 
May. 

c. var. basirotunda Noot. Leid. Bot. Ser. 1 (1975) 

212, pi. 18b. 

Shrub or treelet. Twigs obliquely to patently 
long-pilose. Leaves glabrous to sparsely appres- 
sedly long-pilose beneath, acuminate to caudate 
with rounded to subcordate base and sharply 
glandular dentate to nearly entire margin, elliptic, 
3-1 1 by l 3 4-3 1 2 cm; nerves 6-9 pairs, meeting in a 
looped intramarginal vein; petiole 1-2 mm. 
Flowers in a raceme or panicle to 2 cm, axis 
pilose. Pedicels 0-2(-3) mm. Fruit blue. 

Distr. Malesia: Borneo (Sarawak : Kalabit Up- 
lands). 

Ecol. Montane rain-forest, on humus on sand- 
stone, and on podsolized sand (kerangas), 1000- 
1700 m. Fl. March-April, fr. April, Aug. 

d. var. kinabaluensis (Heine) Noot. Leid. Bot. 
Ser. 1 (1975) 212, pi. 19c. — S. kinabaluensis 
Heine, Mitt. Bot. Staatssamml. Munchen 6 (1953) 
217. 

Shrub or small tree to 4 m. Twigs shortly 
obliquely hairy. Leaves acuminate with cordate 
base and finely glandular-dentate margin, ovate to 
elliptic, l 3 4-4' 2 by l-2 3 4 cm; nerves 4-6 pairs; 
petiole c. ' 2 mm. Flowers in a ± 3-flowered raceme 
to 3 cm, axis with same indument as twigs. Bracts 
3-5 mm, leaf-like, soon caducous. Pedicel l l 2 - 
5 mm. 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu). 

Ecol. Montane rain-forest, also secondary 
forest, and in landslip regrowth, on black or clay 
soils, 1400-2300 m. Fl. Febr., May-Sept., fr. 
March, Aug., Nov. -Dec. 

e. var. mjobcr^ii (Mikk.) Noot. Leid. Bot. Ser. I 
(1975) 212, pi. I8g. — S. mjobergii Mkrk. Sar. 
Mus. J. 3 (1928) 546. - H K . 7. 

Small tree. Twigs patently brown or rusty pilose. 
Leaves (narrowly) elliptic or ovate, acuminate, 
base cordate with 2- 10 mm long lobes, margin 
finely glandular dentate, J 12 by 2 
nerves strongly impressed above, ii 
meeting in a conspicuous looped lotram I 
vein; petiole e. I 1 2 mm. Mowers in ;i P" 
nantly basally branched panicle to tern, il. 
| patently brown or rusty pil. often 



leafiike, and then up to 10 mm. Pedicels 1-5 mm. 
Fruit from green to purple, finally bluish. 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu; 
Sarawak: Mt Murud). 

Ecol. Montane rain-forest, also in secondary 
forest, along hillsides and streams, in Agathis- 
Podocarpus-oak forest, sometimes on blackish 
soil, 1200-2400 m. Fl. Aug.-Nov., fr . Dec-June. 

f. var. pauciHora Noot. Leid. Bot. Ser. 1 (1975) 
213, pi. 18c-d. 

Shrub. Twigs velutinous. Leaves glabrous except 
the appressedly pilose midrib and the recurved 
finely dentate margin underneath, or appressedly 
fine-pilose beneath, acuminate with rounded base, 
elliptic, c. 4 by 2 cm; nerves c. 5-7 pairs, meeting 
in a looped intramarginal vein; petiole with same 
indument as twigs, c. 2 mm. Flowers in a 1-5- 
flowered raceme up to 3 cm ; axis patently pubes- 
cent. Pedicel 0- 1 , mm, but much longer when 
flowers solitary. Fruit blue. 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu; 
Sarawak: Mt Murud). 

Ecol. Montane rain-forest, often mossy, on 
ridges, also in scrub forest, 1700-2570 m. Fl. April, 
July, Oct., fir. April. 

g. var. velutinosa Noot. Leid. Bot. Ser. 1 (1975) 213, 
pi. 19a. 

Treelet to c. 10 m. Twigs velutinous. Leaves 
glabrous above, more or less appressedly pilose 
beneath, especially on the nerves and the sharply 
dentate flat margin, acuminate with rounded base, 
(narrowly) elliptic, 9-12 by 3-4 cm; nerves 7-11 
pairs, meeting in a conspicuous looped intra- 
marginal vein ; petiole 3-4 mm. Flowers in a panicle 
to 3 cm, axis patently pilose. Bracts and the 0-3 mm 
long pedicels with same indument. 

Distr. Malesia: Borneo (Sabah: Mt Kinabalu; 
Sarawak: Kapit area). 

Ecol. Primary and old secondary rain-forest, 
1000-1500 m. Fl. Aug.-Oct. 

34. Symplocos lancifolia S. & Z. Fam. Nat. 2 
(1846) 133; Clarke, Fl. Br. Ind. 3 (1882) 577; 
Brand, Pfl. R. Heft 6 (1901) 41 ; Noot. Leid. Bot. 
Ser. 1 (1975)214, pi. 21a-d, with full synonymy. — 
S. montana Vidal, Rev. PI. Vase. Filip. (1886) 179, 
non Brongn. & Gris, 1866. — S. luzonlensli 
Rolfe.J. Bot. 24(1886) 34X; BRAND, Pfl. R. Hen 6 
(1901) 61; Philip. J. Sc. 3 (1908) Bot. 9; RoLFE, 
Kew Bull. (1912) 157; Brand, Philip. J. Sc. 7(1912) 
Bot. 35; Mikk. En. Philip. 3 (192.1) 300. — S. 
depauperata Mikk. Publ. Cio\. I ah. Philip, n. 29 
(1905) 45; BRAND, Philip. J. Sc. 3 (1908) Mot. 10, 

//;<•/. var. soiiiuh Brand; ibU. i M9i2) Bot 36. 
uul. var. angustissima Brand; Mikk. I n. Philip. 3 
(1923) 298. — S. merrilliana Brand, Philip. J. 
1 1908) Mot. (J . S. bettda Brand, /.<•. B; 
m.kk i n. Philip. 3 (1923) 297; Noot. i rid. Bot 
Ser. I (1975) 133. s. ineonsplcua Brand, Philip 
J. Sc. 4(1909) Bot 110; Mikk. I n. Philip. 3(1923) 
vnboangensii Br ind In I edde, Rep, 14 
(1916) 125; Mikk. I n Philip. 1 (1923) K)3. 
HK. 7. 

hrub I 2 m or tree to 20 m. rwigsappret- 
■edl) to patent!) hairy, toon glabretcent. Leaves 

often sparsely appressedl) fine-hair) beneath, 

acuminate, with cuneate tO iu.nK lounded base 



264 



Flora Malesiana 



[ser. I, vol. 8 2 



and mostly finely glandular dentate or undulate 
margin, (narrowly) ovate, 2-10 by IV2— 4 1 / 4 cm; 
midrib above prominent to slightly sulcate; nerves 
(4— )6— 1 1 pairs, often meeting in a looped intra- 
marginal vein; petiole 1— 3(— 5) mm. Flowers in a 
raceme to 3(-7) cm. Bracts and bracteoles persistent 
but falling in fruit, l U-2 and '/rl'/2 mm respec- 
tively. Pedicel 0-1 mm. Calyx usually sparsely 
appressedly fine short-hairy or pubescent, rarely 
glabrous, divided into ^-I'^rnm long lobes. 
Corolla 2V 2 -4 mm. Stamens 15-40. Disk 5- 
glandular, mostly hairy including the style base. 
Ovary with same indument as calyx or glabrous, 
V2-IV2 mm high. Fruit ellipsoid to globose, 3-5 by 
2-5 mm, the calyx forming a blunt beak on top; 
stone smooth. Seed 1, filling the whole fruit, with 
U-shaped embryo. 

Distr. Continental SE.-E. Asia (N. India, 
Indo-China, China, Hainan, Hong Kong, Ryu Kyu 
Is., Formosa); in Malesia: Philippines (Luzon, 
Mindoro, Panay, Negros, Mindanao). 

Ecol. In a variety of habitats, also in dense 
mossy forest at higher altitude, 400-2500 m. Fl. 
Dec-April, fr. May-Dec. Flowers noted as scent- 
less. 

35. Symplocos lucida (Thunb.) S. & Z. Fl. Jap. 1 
(1835) 55, t. 24, excl. syn. Myrtus laevis; Ohwi, Fl. 
Jap. (1965) 727; Noot. Leid. Bot. Ser. 1 (1975) 217, 
with full synonymy. — Laurus lucida Thunb. Fl. 
Jap. (1784) 174. — Hopea lucida Thunb. Ic. Fl. Jap. 
(1800) t. 4. — S. theaefolia D. Don, Fl. Nepal. 
(1825) 145; Brand, Pfl. R. Heft 6 (1901) 66 
Ctheifolia')\HkL'L.f. Med. Rijksherb. 14(1912)40; 
Back. & Bakh. /. Fl. Java 2 (1965) 205. — 
Dicalyx ciliatus Bl. Bijdr. (1826) 1 1 19. — 5. ciliata 
Miq. Fl. Ind. Bat. 1, 2 (1859) 466; K. & V. Bijdr. 7 
(1900) 155; Brand, Pfl. R. Heft 6 (1901) 65. — 
5. ridleyi King & Gamble, J. As. Soc. Beng. 74, ii 
(1906) 239; Ridl. Fl. Mai. Pen. 2 (1923) 302. — 
5. loheri Brand, Philip. J. Sc. 7 (1912) Bot. 32; 
Merr. En. Philip. 3 (1923) 300. — 5. laeviramulosa 
Elmer, Leafl. Philip. Bot. 7 (1914) 2323; Merr. 
En. Philip. 3 (1923) 300. — Fig. 7. 

Shrub or tree to 20 m, 25 cm 0. Twigs glabrous, 
yellowish green, angular when dry. Leaves coria- 
ceous, glabrous (sometimes quite thin), acute or 
obtuse with cuneate base and entire or glandular 
dentate revolute margin, ± elliptic, 5-12 by 2-A x j 2 
cm; midrib more or less prominent on the upper 
surface, often sulcate towards the base; nerves 
5-15 pairs, prominent on both upper and under- 
surface; petiole 5-15 mm. Flowers in a basally 
branched short dense raceme or condensed spike 
of 17 2 -4 cm; axis puberulous or pubescent. Bracts 
and bracteoles persistent under the fruit, glabrous, 
or sometimes pubescent or puberulous on midrib 
and base, 1-3 mm. Pedicels 0-5 mm. Calyx mostly 
glabrous, nearly divided into 5 lobes, 1-3 mm. 
Corolla 3-5 mm. Stamens 10-70. Disk densely 
hairy. Ovary glabrous, l l 2 -2 mm high; style 
glabrous, or hairy, mostly towards the base. Fruit 
ellipsoid (to rarely nearly orbicular), 1-3-celled, 
5-18 by 4-15 mm, the wider ones with 2 seeds. 
Seeds usually U-shaped with U-shaped embryo, in 
the 3-celled fruits the seeds abortive or (at most) 
V-shaped; the legs of the U are either separated by 
a septum or not. 

Distr. Continental SE.-E. Asia (N. India, N. 



Burma, N. Thailand, Indo-China, China, Hong 
Kong, Hainan, Japan, Ryu Kyu Is., Formosa); 
throughout Malesia, except Borneo, the Moluccas, 
and New Guinea. 

Ecol. High and low mountain forest, elfin forest, 
and mossy forest at higher altitude, also in tjemara 
forest, 1500-3000 m. Fl. (July) Oct.-Nov., fr. July 
(April-Oct.). In habit very much resembling the 
Theaceous Pyrenaria serrata Bl. which grows in 
similar forest. 

Vern. Sumatra: kayu hotir, Asahan; Java: 
djarak lulub, S, djirek, J. 

36. Symplocos maliliensis Noot. Leid. Bot. Ser. 1 
(1975)237. — Fig. 7. 

Tree, 25-30 m, 30-40 cm 0. Twigs glabrous. 
Leaves acuminate, base cuneate, often the very 
base rounded, margin entire, recurved, (narrowly) 
obovate, 15-22 by 4'/ 2 -8 l / 2 cm; nerves 9-14 pairs, 
meeting in a looped intramarginal vein; petiole 
8-15 mm. Flowers in a raceme to 8 cm, axis pubes- 
cent. Bracts and bracteoles caducous, pubescent, 
ovate, 3-4 and 2-3 mm long respectively. Pedicel 
to 2 mm. Calyx glabrous, oblique, 3-4 mm, the 
lobes ovate, 2-3 mm. Corolla c. 6 mm. Stamens c. 
100 or more. Disk shortly pilose. Ovary glabrous, 
1-2 mm high; style with broadly conical shortly 
pilose base, the rest glabrous, c. 5 mm. Fruit 
ellipsoid, 15-20 by 10-12 mm, stone with c. 6 
lengthwise ridges, mostly 2-celled. Seeds not seen. 

Distr. Malesia: Central Celebes (Malili). 

Ecol. Primary high rain-forest, at low altitude, 
c. 200 m. Fl. June-July, fr. Febr., Sept. 

Vern. Lako, kandoa, Tobela lang. 

37. Symplocos wikstroemifolia Hayata, Ic. PI. 
Form. 5 (1915) 119, t. 25b; Mori, Sylvia 5 (1934) 
249; Kaneh. Form. Trees rev. ed. (1936) 602, 
t. 560. — S. microtricha Hand.-Mazz. Beih. Bot. 
Centralbl. 62 B (1943) 17; Noot. Leid. Bot. Ser. 1 
(1975) 239. — Fig. 7. 

Shrub l\/ 2 m, or tree to 20 m. Twigs sometimes 
soon thickened, tapering towards the apex. Leaves 
often only towards the end of the twigs, minutely 
sparsely appressedly fine hairy beneath, acuminate, 
with cuneate base and nearly entire margin, 
(narrowly) elliptic to obovate, 6-1 5V 2 by l 3 / 4 -4 1 / 2 
cm; midrib above prominent or sunken, flat or 
slightly sulcate; nerves 8-10 pairs, joined in an 
intramarginal looped vein 1-3 mm from the 
margin; petiole 3-10 mm. Flowers in an often 
branched spike from the axils of the leaves, the 
lower ones from wood. Bracts and bracteoles soon 
caducous, appressedly pubescent, 1-1 V 4 and 1 mm 
respectively. Flowers <3 or $, probably all flowers 
on one plant alike. Calyx divided into c. 1 mm long 
semi-orbicular or semi-elliptic lobes, glabrous, or 
the outer lobes appressedly fine pubescent. Corolla 
2-3 mm. Stamens 15-20 in $ flowers, 5, alterni- 
petalous, in $ flowers (observed once). Disk pul- 
vinate, glabrous or (minutely) shortly pilose. 
Ovary glabrous or finely appressedly short hairy, 
V 2 mm high in S, l-l' 2 mm in $ flowers; style 
glabrous, 2 mm, with thick, knob-like stigma, but 
aborted in 3 flowers. Fruit ovoid, or slightly con- 
stricted towards the apex, 10-12 by 6-8 mm. Seed 
1, curved, with curved embryo. 

Distr. Continental SE. Asia (Indo-China, 
China, Hainan, Formosa); in Malesia: Malay 



1977] 



Symplocaceae (Nooteboom) 



265 



Peninsula (Pahang: G. Paking, G. Benom, 
Fraser's Hill, G. Tahan). 

Ecol. In hill forest, on mossy or exposed ridges, 
1300-1500 m. Fl. Febr.-March, fr. Oct. In elfin 
forest noted to assume a fastigiate habit. Flowers 
often in part ramiflorous. 



38. Symplocos multibracteata Noot. Leid. Bot. Ser. 
1 (1975) 241. — Fig. 19. 

Small shrub, i 4 -l' 4 m, or treelet to 4 m. Twigs 
densely appressedly to patently (softly) pilose to 
nearly glabrous. Leaves acuminate with rounded to 
cordate base and glandular denticulate to dentate 
margin, elliptic to ovate, 5-14 by ZVz-SVaCm; 
nerves 7-13 pairs, meeting in a looped intramargi- 
nal vein; petiole 2-10 mm. Flowers in a reduced 
spike of at most 2 cm, usually only 1 (subterminal) 
flower left, sometimes another flower present in 
bud, axis glabrous. Bracts many, appressedly 
pubescent, 4-8 cm. Calyx divided into the appres- 
sedly pubescent 3-5 mm long lobes. Corolla 5-8 
mm. Stamens 80 to more than 150. Disk softly 
pilose. Ovary glabrous, 2-3 mm high; style gla- 
brous, to 2 1 2 mm long. Fruit obliquely ovoid to 
ellipsoid to spindle-shaped, 17-22 by 8-10 mm. 
Seed 1, filling the whole stone, with the embryo 
straight or slightly curved. 

Distr. Malesia: East New Guinea (W. & E. 
Highlands). 



Ecol. Montane rain-forest and depleted 
Castanopsis-Nothofagus forest, 2000-2300 m. Fl. 
July, Sept.,//-. Aug., Jan. 

Vern. Chandujant, Wabag, Enga lang. 

39. Symplocos nivea Brand, Pfl. R. Heft 6 (1901) 
36; K. & G. J. As. Soc. Beng. 74, ii (1906) 234; 
Ridl. Fl. Mai. Pen. 2 (1923) 300; Noot. Leid. Bot. 
Ser. 1 (1975) 241. 

Tree to 18 m. Twigs glabrous. Leaves acuminate 
with cuneate, attenuate base and entire to obscurely 
undulate-crenate margin, glabrous, (narrowly) 
elliptic, 7-11 by 2-4'/ 2 cm; nerves 5-8 pairs, 
meeting in an intramarginal vein 2-5 mm from the 
margin; petiole 7-10 mm. Flowers in a panicle of 
racemes, axis villous. Bracts and bracteoles 
glabrous, soon caducous, 2V 2 -3 and c. 2 l ! 2 mm 
long respectively. Pedicel pubescent, to 5 mm long. 
Calyx glabrous, 2'/ 2 -3 mm, the lobes 1-2 mm long, 
becoming longer by tearing apart. Corolla c. 5 mm. 
Stamens more than 100. Disk 5-glandular, with the 
broadly conical style base soft hairy. Ovary 
glabrous, c. 1 mm high; style glabrous, c. 5 mm. 
Fruit not known. 

Distr. Malesia: Malay Peninsula (Penang, 
Johore), 2 collections. 

Ecol. Hill rain-forest. 

Note. Closely allied to 5. pyriflora Ridl., 
differing in the number of nerves and with shorter 
corolla. May in future prove to be conspecific. 




1 Symplocot mtitrtbn* xeata N001 1 Leafand flower, nat. size, b. deflorated Rower, ihowina s lobed 
iisk, c. fruit, <i endocarp, e. fruit in CS f all 2 (a-b Hoooland 5882, e t ii<xm,i and 5«k7). 



266 



Flora Malesiana 



[ser. I, vol. 8 2 



40. Symplocos obovatifolia Merr. Philip. J. Sc. 12 
(1917) Bot. 290; En. Philip. 3 (1923) 300; Noot. 
Leid. Bot. Ser. 1 (1975) 242. — Fig. 7. 

Twigs glabrous. Leaves glabrous, rounded or 
shortly acuminate with cuneate, attenuate base and 
entire or glandular denticulate apex, obovate, 
7 1 / 2 — 1 1 by 3'/2-6 cm; nerves 7-9 pairs, meeting in a 
looped intramarginal vein; petiole 7-12 mm. 
Flowers in a fascicle or short spike to P/2 cm, axis 
glabrous. Bracts and bracteoles glabrous, per- 
sistent, 2-3 mm. Only fruits seen. Calyx 3-lobed, 
glabrous, elliptic, rounded, c. 2 mm. Disk glabrous, 
style base shortly pilose. Fruit (obliquely) ellipsoid, 
c. 11 by 5 mm, the persistent calyx not included; 
stone smooth, 3-celled. Seed 1 in each cell, straight 
with straight embryo. 

Distr. Malesia: Philippines (Luzon, Mt Umin- 
gan, Nueva Ecija), 2 collections. 

Ecol. Hill rain-forest. Fr. Aug.-Sept. 

41. Symplocos odoratissima (Bl.) Choisy ex Zoll. 
Syst. Verz. 2 (1854) 136; Miq. Fl. Ind. Bat. 1, 2 
(1859) 468; Gurke in E. & P. Nat. Pfl. Fam. 4, 1 
(1891) 170; K. & V. Bijdr. 7 (1900) 148, incl. var. 
aluminosa K. & V. I.e. 150; Brand, Pfl. R. Heft 6 
(1901) 35, incl. var. divaricata Brand; K. & G. J. 
As. Soc. Beng. 74, ii (1906) 233; Koord. Atlas 2 
(1914) t. 382; Ridl. Fl. Mai. Pen. 2 (1923) 299; 
S. Moore, J. Bot. 63 (1925) Suppl. 65, incl. var. 
leptocarpa S. Moore; Heyne, Nutt. PI. (1927) 
1263; Burk. Diet. 2 (1935) 112; Back. & Bakh./. 
Fl. Java 2 (1965) 205; Noot. Leid. Bot. Ser. 1 
(1975) 245. — Dicalyx odoratissimus Bl. Bijdr. 
(1826) 1116. — Eugeniodes odoratissima O. K. Rev. 
Gen. PI. 2(1891)975. 

For further synonyms see under the varieties. 

Tree (shrub) to 30 m high and 50 cm 0. Twigs 
glabrous or tomentellous to tomentose or pubes- 
cent. Leaves glabrous or pubescent beneath, 
especially on midrib and nerves, with blunt, usually 
acuminate apex, acute to rarely rounded base and 
entire or mostly crenulate or dentate margin, 
(narrowly) elliptic to obovate, 7-20(-40) by 
(2V2-)5-10(-20) cm; nerves 5-13(-16) pairs; 
petiole stout, 1-5 cm. Flowers mostly many in a 
5-30 cm long panicle which is sometimes only 
branched towards the base, the axes rusty tomen- 
tellous. Bracts at the base of the 3-7 mm long 
pedicel, 3-5 mm, bracteoles directly under each 
flower, both tomentellous on both surfaces, 
caducous. Calyx tomentellous, the lobes blunt, 
V2-IV2 mm - Corolla usually tomentellous, at least 
in bud, rarely nearly glabrous, 5-8 mm. Stamens 
more than 100. Disk hairy with 5 conspicuous 
glands. Ovary with same indument as calyx, 
1 V2-2V2 mm high ; style pilose towards the conical 
base, about as long as the corolla. Fruit glabrous or 
tomentellous, (obliquely) ovoid (or rarely narrowly 
flask-shaped, pear-shaped or globular), more or less 
narrowed towards the apex, 8-25 by 5-20 mm ; 
stone with c. 5(— 10) ridges. Seeds curved, with 
curved embryo. 

Distr. Throughout Malesia, except New Guinea. 

key to the varieties 

1. Twigs mostly glabrous. Leaves 7-23 cm long. 
Fruit 8-1 5(-20) by 5-10 mm. 

a. var. odoratissima 



1. Twigs mostly patently pilose, pubescent or 
tomentose. Leaves 15-27 cm long. Fruit 17-25 
by 1 2-20 mm b. var. wenzelii 

a. var. odoratissima. — Cf. Noot. Leid. Bot. Ser. 1 
(1975) 247. — Dicalyx odoratissimus Bl. Bijdr. 
(1826) 1116. — Dicalyx aluminosus {non Lour.) 
Bl. I.e. 1117, p.p. — S. ciliata Presl, Rel. Haenk. 
2 (1831) 61; F.-Vill. Nov. App. (1880) 127. — 
S. patens Presl, Rel. Haenk. 2 (1831) 61 ; F.-Vill. 
Nov. App. (1880) 127; Brand, Pfl. R. Heft 6 (1901) 
34, incl. var. ciliata Brand, I.e. 35 ; Philip. J. Sc. 3 
(1908) Bot. 4, incl.f. ciliata Brand, I.e. 5 etf. elmeri 
Brand, I.e. 4; Merr. En. Philip. 3 (1923) 301. — 
S. repandula Miq. Fl. Ind. Bat. Suppl. 1 (1861) 474. 

— S. racemosa (non Roxb.) F.-Vill. Nov. App. 
(1880) 127. — S. spicata (non Roxb.) F.-Vill. I.e. 
127; Vidal, Sinopsis Atlas (1883) t. 64. — S. vil- 
larii Vidal, Rev. PI. Vase. Filip. (1886) 178, excl. 
syn. Guettarda polyandra Blanco, nom. illeg. — 
S. pseudospicata Vidal, I.e. 179. — Pygeum grandi- 
florum King, J. As. Soc. Beng. 66, ii (1897) 228; 
cf. Kalkman, Blumea 13 (1965) 107. — S. alumi- 
nosa Brand, Pfl. R. Heft 6 (1901) 35. — S. poly- 
andra sens. Brand, I.e. 36, quoad descr. et syn. 
Vidal. — S. floridissima Brand, I.e. 35; Philip. J. 
Sc. 3 (1908) Bot. 5; ibid., 7 (1912) Bot. 32; Merr. 
En. Philip. 3 (1923) 298. — S. elmeri Brand in 
Perkins, Fragm. Fl. Philip. (1904) 36. — S. pul- 
verulenta King & Gamble, J. As. Soc. Beng. 74, ii 
(1906) 234; Ridl. Fl. Mai. Pen. 2 (1923) 300; Burk. 
Diet. 2 (1935) 112. — S. floridissima Brand var. 
serrata Brand, Philip. J. Sc. 4 (1909) Bot. 108. — 
S. pulgarensis Elmer, Leafl. Philip. Bot. 5 (1913) 
1841 ; Merr. En. Philip. 3 (1923) 302. — S. apoen- 
sis Elmer, Leafl. Philip. Bot. 7 (1914) 2319; Merr. 
En. Philip. 3 (1923) 297. — S. megabotrys Merr. 
Philip. J. Sc. 9 (1914) Bot. 383 ; En. Philip. 3 (1923) 
300. — S. dagamensis Brand in Fedde, Rep. 14 
(1916) 324; Merr. En. Philip. 3 (1923) 298. — 
S. salix Brand in Fedde, Rep. 14 (1916) 325; 
Merr. En. Philip. 3 (1923) 302. — S. acuminatis- 
sima Merr. Philip. J. Sc. 11 (June 1916) Bot. 31; 
En. Philip. 3 (1923) 296. — Pygeum viride Baker 
/. J. Bot. 62 (1924) Suppl. 34; cf. Kalkman, 
Blumea 13 (1965) 107. — S. bulusanensis Elmer, 
Leafl. Philip. Bot. 10 (1939) 3792, nom. illeg., angl. 

— S. verdifolia Elmer, I.e. 3793, nom. illeg., angl. 
—Fig. 7. 

Tree up to 30 m, 50 cm 0. Twigs, petioles and 
underside of leaves mostly glabrous, sometimes 
however tomentellous, tomentose, or pubescent. 
Leaves 7-23 by 2-12 cm, in watersprouts up to 
40 cm. Fruit with thin, fleshy mesocarp, 8-15 by 
5-10 mm, ovoid, or up to 20 mm, flask-shaped. 
The stone with low ridges. 

Distr. Throughout Malesia, except New 
Guinea. 

Ecol. Primary and secondary rain-forest, not 
rarely along river-banks, on sandy river alluvium, 
in Borneo also on brown sandy soils, black soils, 
loam and sandstone, from sea-level to 2500 m. Fl. 
Febr. -Nov., fr. Aug.-March. Flowers are noted to 
be fragrant. 

Uses. Dayak people extract salt from wood ash. 

As in other species the bark is used for dyeing 
purposes and Heyne I.e. even says that for the 
purpose of obtaining bark and leaves the species is 
planted by the Sundanese at Tjiamis. 



1977] 



Symplocaceae (Nooteboom) 



267 



The tree is mainly useful for the inner bark which 
is commonly sold in the medicinal market in West 
Java as kayu or kulit seriawan. Decoctions are used 
against sprue-like diseases; also pounded bark is 
applied to the gums and young leaves are some- 
times eaten or applied externally on mouth and 
nose. Obat seriawan is even officially recognized in 
the Dutch pharmacopeia. 

Vern. Sumatra: sarigintung, Karo, tjirupago 
uding, Simalur; Java: ki njatu, ki sariawan, ki 
siriawan, S; Borneo: lisang, Kinabatangan, Dusun 
lang., margaram, Sangkulirang I.; Bali: udu; 
Talaud: labah. 

b. var. wenzelii (Merr.) Noot. Leid. Bot. Ser. 1 
(1975) 248. — S. wenzelii Merr. Philip. J. Sc. 10 
(1915) Bot. 282; En. Philip. 3 (1923) 302. — 
S. trichophlebia Merr. Un. Cal. Publ. Bot. 15 
(1929) 248. — Fig. 7. 

Tree up to 26 m, 50 cm 0. Twigs usually patently 
pilose, pubescent or tomentose. Leaves mostly 
densely pubescent, 15-27 by 12-20 cm; ridges on 
the stone up to 4 mm. 

Distr. Malesia: Borneo (Sarawak and Kaliman- 
tan), Philippines (Leyte, once). 

Ecol. Primary and secondary rain-forest in the 
lowland and hills in a variety of conditions: sandy 
ridges and slopes, calcareous loam, dark red soil, 
and black soil, near streams. Fl. (March) June- 
Dec., fir. (Febr.-May) July. Obviously mature 
fruits are often noted pale green or white. 

Note. Size and shape of leaves are very variable 
in S. odoratissima; var. wenzelii possesses the larger 
and most hairy leaves. The flowers are exactly 
matching those of var. odoratissima and with 
collections without fruit it is not always possible to 
decide to which variety they belong. 

42. Symplocos ophirensis Clarke, Fl. Br. Ind. 3 
(1882) 579; K. & G. J. As. Soc. Beng. 74, ii (1906) 
243;Ridl. Fl. Mai. Pen. 2 (1923) 305; Noot. Leid. 
Bot. Ser. 1 (1975) 249, f. 4a-e. — Eugeniodes 
ophirense O. K. Rev. Gen. PI. 2 (1891) 975. — 
Fig. 4a-e. 

For further synonyms see under the infraspecific 
taxa. 

Shrub or tree to 18 m high and 50 cm 0. Twigs 
glabrous, or sometimes the youngest parts appres- 
sedly pubescent. Leaves glabrous, except some- 
times the very youngest, cuneate or rounded to 
acuminate, with cuneate base and entire, glandular 
crenulate to denticulate or serrate margin, elliptic 
to ovate or obovate, 5-22 by l'/ 2 -7cm; nerves 
4—1 3(-l 6> pairs, anastomosing or meeting in an 
intramarginal vein; petiole 2-10(-20) mm. Flowers 
in a short raceme, a 3-5-branched panicle of 
racemes or a spike of I -3(-6 in Sumatra) cm, 
rarely only 1-3 flowers together; axis apprcssecll> 
pubescent to minutely puberulous or nearly gla- 
brous. Bracts and bracteoles caducous or persistent, 
with same indumcnt as axis, '/i •' .' fl mm anU " 
slightly shorter than that respectively. Calyx with 
same indumcnt as ovary or less hairy, '/i I (2'/i ^ 
in ssp. cumingianu var. paih\[>li\lla) mm long. 
Corolla 2 5 mm. Stamens 20 60, but more than 75 
invar, pat hyphylla. Disk glabrous to shoitly pil".e. 
5-glandular. Ovary mostly with same inilument as 
inflorescence axis, or densely appressedly pubes 
cent, rarely glabrous, c. l'/j rnrn high (2 1 /, mm m 



var. pachyphylla); style glabrous to pilose, 3-5(-8) 
mm. Fruit ampulliform, with long neck, to ovoid, 
rarely ellipsoid or cylindrical; stone with coarse 
surface, low lengthwise ridges, or high, interrupted 
ridges and then with hollow base, filled with fleshy 
mesocarp. Seed 1, embryo obscurely S-shaped, 
curved with an angle of c. 90 c ± halfway its length, 
or twice screw-like curved. 

Distr. Malesia: Central West Sumatra (incl. 
Lingga Is.), Malay Peninsula, Borneo, Celebes, and 
throughout the Philippines. 

KEY TO THE INFRASPECIFIC TAXA 

1. Twigs densely appressedly pubescent or tomen- 
tose . 1. ssp. ophirensis b. var. densireticulata 
1. Twigs glabrous or sparsely fine-hairy. 
2. Fruit ampulliform, with long neck. Ovary 
P^rnm high. Calyx lobes c. '/j mm long. 
Corolla 2-3 mm. Disk globose or annular, 

shortly pilose 2. ssp. perakensis 

3. Terminal buds glabrous. Secondary veins 
forming a rather coarse reticulation with the 
slightly less prominent veins. Inflorescence a 
many-flowered panicle of racemes, 1-4 cm 
long. Style shortly pilose for its whole length. 
c. var. perakensis 
3. Terminal buds glabrous. Secondary veins 
prominent, forming a fine reticulation with the 
faintly prominent tertiary veins. Inflorescence 
a 1-3-flowered raceme, up to 1 cm. Style 
pilose only towards its base. 

d. var. lingaensis 

3. Terminal buds pubescent. Secondary and 
tertiary veins much prominent, forming a fine 
reticulation with the often also prominent 
quaternary veins. Style glabrous. 

e. var. sumatrana 

2. Fruit ovoid, ellipsoid, or cylindrical. Ovary 

Vl 2 (-2 l l 2 )mm high. Calyx lobes Vr-K-lVa) 

mm long. Corolla 3-5 mm. Disk 5-glandular, 

glabrous or sparsely hairy. 

4. Fruit ovoid to cylindrical; stone with shallow 
lengthwise grooves. Seed ovoid, with small, 
nearly straight embryo. Disk sparsely hairy. 
Reticulation beneath very dense. 

1. ssp. ophirensis a. var. ophirensis 

4. Fruit ovoid, ellipsoid, or rarely cylindrical; 

stone with high, interrupted ridges. Seeds 

ovoid to horse-shoe-shaped, embryo curved, 

or twice screw-like curved. Disk glabrous, 

rarely with some hairs. Reticulation beneath 

either very fine or coarse 3. ssp. cumingianu 

5. Inflorescence a raceme. Ovaiy 1 ' - mm. 

Calyx lobes ' a I mm. Corolla 3 4 1 , mm. 

Stamens 20 60 . . . f. tar. cumingiana 

5. Inflorescence ■ spike ()\.u> 2' , mm. ( 'al>\ 

lobes e. I' 2 mm. CoroOa c. 3 mm. stamens 
more than 75 ... . g. var. pachyphylla 

I ophirensis. — Cf. N<h>i I eul. BOi Sei I 

(1975) 252. s ophlrensit i i mucb, I L Br. Ind. 3 

(IKK2) 579. 

I .'i the description see the ipe 

a. var. ophirensis. I- in. 7. 

Shnib I ' i m, oi small tiee I" 6 OL I ca\< 

mm. lie "i rounded, (narrowly) elliptic. S 1 ; 9* : 

( II 1 ,) In I' B; nerves J 6 pans. 



268 



Flora Malesiana 



[ser. I, vol. 8 : 



meeting in a looped intramarginal vein; petiole 
only with faint ridges towards the blade. Racemes 
up to 10 mm, from the axils of the upper leaves or 
from wood. Bracts caducous or persistent. Pedicels 
1-3 mm. Calyx lobes l l 2 -\ mm. Corolla 3-5 mm. 
Stamens 25-60. Disk usually sparsely hairy. Ovary 
lV 2 mmhigh; style glabrous, 3 1 / 2 -5 mm. Fruit 
ovoid to cylindrical, 6-12 by 4-5 mm. Seed ovoid, 
with small, nearly straight embryo. 

Distr. Malesia: Malay Peninsula (Perak, 
Selangor, Johore). 

Ecol. Montane forest, bush-like, on granite, 
1200-1500 m. Fl. July-Sept., fr. Aug. Young leaves 
black purple. 

b. var. densireticulata Noot. Leid. Bot. Ser. 1 (1975) 
252. 

Small, bushy treelet, 2-4 m. Twigs (appressedly) 
pubescent to tomentose. Leaves cuneate to acumi- 
nate with cuneate to cordate base, 3V2-1 1 by 
l 1 / 2 -4 1 /2cm; nerves 6-9 pairs, anastomosing or 
meeting in an intramarginal vein; petiole 2-9 mm. 
Flowers in a short raceme to c. 3 cm ; axis pubes- 
cent. Bracts and bracteoles pubescent, soon 
caducous, 2 and 1 mm long respectively. Calyx 
pubescent, l-l\/ 2 mm long, the lobes ± triangu- 
lar, c. 1 mm. Corolla 2-2V 2 mm. Stamens delicate, 

c. 40. Disk inconspicuous, pilose. Ovary pubescent, 

1 mm high; style glabrous, c. 2 mm. Fruit pubes- 
cent, ellipsoid, 5-8 by 4-5 mm; stone smooth. 
Seed not seen. 

Distr. Malesia: Malay Peninsula (Pahang: 
Cameron Highlands) and S. Celebes, in both areas 

2 collections each. 

Ecol. Montane forest, 1400-2500 m. Fl. Sept. 

2. ssp. perakensis (K. & G.) Noot. Leid. Bot. 
Ser. 1 (1975) 254. — S. perakensis King & Gamble, 
J. As. Soc. Beng. 74, ii (1906) 241 ; Ridl. Fl. Mai. 
Pen. 2 (1923) 304; Burk. Diet. (1935) 2114. — 
S. caudata (non Wall, ex G. Don) Ridl. Fl. Mai. 
Pen. 2 (1923) 304. 

Tree to 18 m high and 50 cm 0. Leaves faintly 
acuminate to caudate with cuneate base, (narrowly) 
elliptic, 5-12 by 2-4\' 2 cm; nerves 4-7 pairs, except 
in var. sumatrana meeting in a looped intramarginal 
vein; petiole 3-9 mm, not winged. Flowers in a 
(basally) 3-5-branched very slender panicle of 
racemes, a raceme, or in var. lingaensis only 1-3 
flowers in each inflorescence. Bracts and bracteoles 
persistent, minute. Pedicels 1-4 mm. Calyx divided 
into semiorbicular c. l /j mm long lobes. Corolla 
2-3 mm. Stamens 30-50. Disk shortly pilose. Ovary 
l /r-l mm high; style pilose to glabrous. Fruit 
ampulliform, c. 1 by 5 mm, with long beak; stone 
with coarse surface, the inner wall of the stone 
following the grooved surface of the deeply rumi- 
nate cerebrum-like seed; embryo curved with an 
angle of not yet 90°. 

Distr. Malesia: Sumatra, Malay Peninsula, and 
the Philippines. 

c. var. perakensis. — Cf. Noot. Leid. Bot. Ser. 1 
(1975) 255, f. 4a-c. — 5. fragrans Elmer, Leafl. 
Philip. Bot. 2 (1908) 508; Brand, Philip. J. Sc. 7 
(1912) Bot. 33; Merr. En. Philip. 3 (1923) 299. — 
Fig. 4a-c, 7. 

Leaves 5-11 by 2-4V 2 cm; petiole 3-6 mm. 
Flowers in a many-flowered panicle of racemes of 



1-4 cm. Calyx and ovary appressedly pubescent; 
style shortly pilose for its whole length. 

Distr. Malesia: Malay Peninsula and the Philip- 
pines (Negros, once). 

Ecol. Primary lowland and montane forest, 
hillsides, bamboo forest, 60-1500 m. Fl. April- 
July, Sept.,/r. Nov. 

d. var. lingaensis Noot. Leid. Bot. Ser. 1 (1975) 255. 
Leaves narrowly elliptic with caudate apex, 

7-12 by 2-3 1 / 2 cm; petiole c. 5 mm. Flowers in a 
1-3-flowered raceme to 1 cm. Calyx and ovary 
minutely puberulous; style pilose only towards its 
base. Fruit unknown. 

Distr. Malesia: Sumatra (Lingga Arch.). Only 
known from the type. 

e. var. sumatrana Noot. Leid. Bot. Ser. 1 (1975) 
256. 

Leaves faintly acuminate, narrowly elliptic, 
6-10 by 2-3 cm; nerves 4-5 pairs; petiole 4-9 mm. 
Flowers in a lax panicle or raceme of lV 2 -6cm. 
Calyx and ovary minutely appressedly pubescent; 
style glabrous. Fruit not known. 

Distr. Malesia: Central West Sumatra. 

Ecol. Montane forest, 900-1300 m. 

3. ssp. cumingiana (Brand) Noot. Leid. Bot. Ser. 
1 (1975) 253. — S. cumingiana Brand, Pfl. R. Heft 
6(1901) 58. 

Shrub or small tree to 6 m. Leaves ± elliptic, 
6-22 by 3-7 cm; nerves 6— 13(— 16) pairs; petiole 
l-10(-20) mm, narrowly winged, except to its very 
base. Flowers in a 3(-5) cm long often branched 
raceme or spike. Pedicels 0-3 mm. Calyx V 2 -l 
(-l 1 / 2 )mm long. Corolla 3-5 mm. Disk 5-glan- 
dular, glabrous, rarely with some hairs. Ovary 
1V 2 (-2V 2 ) mm high. Fruit ovoid, ellipsoid or rarely 
cylindrical, 5-12 by 3-8 mm; stone with high, 
interrupted ridges which often protrude from the 
base, enclosing some fleshy mesocarp. Seed ovoid 
to horse-shoe-shaped, embryo curved with an angle 
of about 90^ to twice screw-like curved. 

Distr. Malesia: Borneo, Philippines and 
Celebes. 

f. var. cumingiana. — Cf. Noot. Leid. Bot. Ser. 1 
(1975) 253, f. 4d-e, pi. 20a-e. — S. cumingiana 
Brand, Pfl. R. Heft 6 (1901) 58; Philip. J. Sc. 3 
(1908) Bot. 8; ibid. 7 (1912) Bot. 34; Merr. En. 
Philip. 3 (1923) 297 ; H. Heine, Pfl. Samml. Clemens 
Kinabalu (1953) 87. — S. curtiflora Elmer, Leafl. 
Philip. Bot. 2 (1908) 509; Merr. En. Philip. 3 
(1923) 298. — S. angularis Elmer, Leafl. Philip. 
Bot. 2 (1908) 510. — S.purpurascens Brand, Philip. 
J. Sc. 7 (1912) Bot. 33; Merr. En. Philip. 3 (1923) 
302. — S. minutiflora Elmer, Leafl. Philip. Bot. 7 
(1914) 2320; Merr. En. Philip. 3 (1923) 300. — 
S. agusanensis Elmer, Leafl. Philip. Bot. 7 (1914) 
2321. — S. elliptifolia Merr. Philip. J. Sc. 12 
(1917) Bot. 292; En. Philip. 3 (1923) 298. — S. 
brachybotrys Merr. Philip. J. Sc. 14 (1919) 447, 
non Merr. 1917; En. Philip. 3 (1923) 297. — 
S. ilocana Merr. Philip. J. Sc. 35 (1928) 7. — Fig. 
4d-e, 7. 

Shrub lV 2 m or small tree to 12 m, once even 
30 m and 50 cm 0. Leaves ± elliptic, 6-18 by 
3-7 cm; nerves 6— 13(— 16) pairs, usually meeting 
in a looped intramarginal vein; petiole 3-10(-15) 



1977] 



Symplocaceae (Nooteboom) 



269 



mm. Racemes to 3(-5) cm long. Bracts and brac- 
teoles usually very small, caducous or persistent. 
Pedicels 1-3 mm. Calyx 1 / 2 — 1 mm, pubescent. 
Corolla 3-4 1 , mm. Stamens 20-60. Ovary l'/ 2 mm 
high. Fruit 5-12 by 3-7 mm, ripe purple-blue. 

Distr. Malesia: Borneo, Philippines, Celebes. 

Ecol. Mostly in the mountain forest, on hillsides 
in oak-Fodocarpus forest, largely between 1000 and 
3000 m, but on Mt Kinabalu once found as high as 
3700 m (sterile), and once collected in lowland 
Dipterocarp forest at 300 m in the Sierra Madre 
Mts (Luzon), a very common species in the 
Philippines. Flowers (once) noted to be faintly 
fragrant. Fl. May-Dec, fr. March-Oct. 

g. var. pachvphvlla (Merr.) Noot. Leid. Bot. Ser. 1 
(1975) 254. — S. pachyphylla Merr. Philip. J. Sc. 
10 (1915) Bot. 283. — Fig. 7. 

Small tree, 6 m. Leaves 10-20 by 6-8 1 / 2 cm; 
nerves c. 10 pairs; petiole 10-20 mm. Flowers in a 
spike. Bracts and bracteoles appressedly pubescent, 
2Vi and 3 mm long respectively. Calyx densely 
appressedly pubescent, divided into c. 1 l /a mrn ' on 8 
lobes. Corolla c. 5 mm. Stamens more than 75, up 
to 9 mm. Disk glabrous, 5-glandular. Ovary 
glabrous, 2 1 4 mm high ; style glabrous, c. 8 mm. 
Fruit ovoid, c. 10 by 6-8 mm, the stone as in var. 
cumingiana, but several ridges totally lacking in the 
upper half, c. 1 by 5-6 mm. Seed ovoid or curved, 
and then as the embryo with an angle of about 90° 
beneath the middle. 

Distr. Malesia: Philippines (Leyte and Minda- 
nao), 2 collections. 

Ecol. Hill forest, c. 500 m. Fl. Sept. 

43. Symplocos paucistaminea F.v.M. & F. M. 
Bailey, 3rd Suppl. Syn. Queensl. Fl. (1890) 46; 
F. M. Bailey, Queensl. Fl. 3 (1900) 967; Noot. 
Leid. Bot. Ser. 1 (1975) 262. — Fig. 7. 

Tree 18 m, 45 cm 0. Twigs densely spreadingly 
pubescent to tomentose. Leaves acuminate, with 
acute to rounded base and dentate margin, sparsely 
pubescent above and beneath, elliptic to obovate, 
8-20 by 3-8 cm; nerves 7-12 pairs, meeting in a 
looped intramarginal vein; petiole 5-10 mm. 
Flowers in a basally branched spike to 5 cm long, 
becoming longer in fruit ; axis sparsely brown hairy. 
Bracts and bracteoles persistent, spreadingly 
hairy, c. 2 and c. l'/ 2 mm respectively. Calyx 
divided into glabrous c. 1 mm long lobes, the lobes 
tomentose. Corolla c. 2 l / 2 mm - Stamens c. 10 to 60. 
Disk glabrous or pilose. Ovary glabrous, c . 3 / 4 mm 
high; style glabrous, c. l'/^rnm. Fruit ampulliform, 
c. 6 by 4 mm, stone ampulliform with globose, 
lengthwise grooved belly and narrow cylindrical 
neck, I -celled. Seed 1, filling the whole stone, with 
the embryo twice curved. 

Distr. Queensland and Malesia (New Guinea: 
Milne Bay Distr.: Mt Suckling, two collections). 

Ecol. Lowland rain-forest at 360 m. Fl. July. 

44. S\mplocos polyandra (BLANCO) BltAND, I'll 
K Hefto (1901) 436, quoad syn. Blanco, exel desa 
etitirp.;Man Sp. Blanc. (1918) 304; En Philip. 3 
(1923) 301; sin-, Hull. Bot. Card Btzg ill, 12 
(1932) 170, f. 5; Noot. Leid. Bot. Ser, I (1975) 
264. - Guettarda polyandra Hi kn* o, Fl. 1 [lip, ed. 
2 (1845) 500; ed. i ( IK7«>> 126. < arlea oblongi- 



folia Presl, Epim. Bot. (1851) 216. — Baranda 
angatensis Llanos, Mem. Acad. Cienc. Madrid 3, 
2 (1857) 502. — S. oblongifolia Rolfe, J. Bot. 23 
(1885) 214; Vidal, Phan. Cuming. Philip. (1885) 
124; Rev. PI. Vase. Filip. (1886) 178; Brand, Pfl. 
R. Heft 6 (1901) 55; Hall./. Beih. Bot. Centralbl. 
39 B (1921) 94. — S. superba Brand, Pfl. R. Heft 6 
(1901) 55. — Fig. 7. 

Tree up to 30 m, 50 cm 0, rarely a shrub. Bark 
dark, cracked. Twigs puberulous, glabrescent, 
tapering off towards the apex, thick, at least 5 mm 
beneath the leaves and there usually with many 
pulvinate leaf-scars. Leaves crowded towards the 
end of the twigs, rounded or cuneate-obtuse at the 
apex, with cuneate, attenuate base and entire, 
revolute margin, glabrous (except in innovations 
and then puberulous), narrowly elliptic to obovate, 
9-22 by 2'/ 2 -7(-9) cm; nerves 11-15 pairs; petiole 
2-4 cm. Many spikes from old wood beneath the 
leaves, axis densely rusty appressedly puberulous, 
glabrescent, 4-15 cm long. Bracts and bracteoles 
with same indument, persistent under the fruit, 
lV2-2mm long. Calyx with same indument, 
becoming glabrous towards the apex, 2-3 mm, the 
lobes c. 2 mm long. Corolla 8-10 mm. Stamens 50 
to more than 100. Disk glabrous, annular, and 
then surrounding a lower, rarely shortly pilose 
receptacle, or low pulvinate, only surrounding the 
glabrous, 7-9 mm long style. Ovary with same 
indument as calyx, c. 2 mm high. Fruit ellipsoid, 
c. 10 by 7 mm in vivo; stone rather smooth, with 
few shallow lengthwise grooves, 8-10 by 4-5 mm 
(s.s. the whole fruit as big as the stone), 3-celled. 
Seed 1 in each cell with straight embryo. 

Distr. Malesia: Borneo and adjacent islands 
(Natuna, Banka, Billiton, Karimata, St. Barbe), 
Philippines (throughout), and SW. Celebes 
(Makassar: Baleh Angien, once). 

Ecol. Secondary and primary forest, almost 
always on sandstone, granite, kerangas, sandy 
flats, more rarely on sandy loam, at low altitude, 
below c. 300 m, once found in montane forest 
(Luzon: Sierra Madre) at 1000-1 100m in low, 
mixed, primary rain-forest (Jacobs 7840). Fl. 
Sept.-March, fr. Febr.-June (July-Oct.). The 
flowers are faintly fragrant, especially at night. 

Vern. Bungur, dutat, Banka; sudjeng, Naiuna; 
Borneo: merbryot, Sarawak, hcluno-beluno. salum- 
buno, temasuk jantan, Sandakan; Philippines: 
ditdman, rapo-rdpo, Tag., balakbdk, balakbakan, 
bangkuned, mankdnai, P.Bis., buH-buli, malabaU, 
ribuli, Pang., dilangi-bdka, Sbl. 

45. Symplocos pulvinata Noot. Leid. Hoi Ser. I 
(1975)269. — Fig. 7. 

Sparsely foliaged tree, 12 18 m high. Twigs thick, 
;ii least 5 mm. Leaves coriaceous, glabrous, acute 
in Faintly acuminate with cuneate base ami glan- 
dular crenate oi dentate margin, obovate, 12 21 by 

4*/j 10'. .cm; nerves K 12 pans; petiole stout, 

I 1 , 2' . Lin. Spike glabrous, e. 1 cm. Bra< 
bracteoles probably persistent, glabrous, 3 ~> ami 
, i nun long respective!) (olda Bowers often 
Fallen tncludmfl bracts .mil bracteoles, leaving 
conspicuous pulvinate lighl coloured scars on the 
(i, nk axis) ( "/i i glabrous, i I mm. divided In 5. 
mm long i"i * in some (lowers < orolla ami 

si. miens absent 01 obsolete, in othei llowets 






270 



Flora Malesiana 



[ser. I, vol. 8 2 



corolla 5 mm, 3(-4)-lobed and stamens 20-35. 
Disk glabrous. Ovary glabrous, oblique, 1-1 V2 mm 
at one, c. 2 mm at the other side; style glabrous, 
6 mm. Fruit ovoid, deeply violet, c. 13 by 6-8 mm; 
stone with rather high lengthwise ridges in the 
basal and low ridges in the apical half, in the middle 
a deep transverse groove, 1-celled. Seed 1, 
uncinately curved towards the base with curved 
embryo. 

Distr. Malesia: East New Guinea (Koitaki and 
Normanby I.), 2 collections. 

Ecol. Under open canopy of tall forest, 450- 
825 m. Fl. Febr. 

46. Symplocos pyriflora Ridl. J. Fed. Mai. St. 
Mus. 6 (1915) 159; Fl. Mai. Pen. 2 (1923) 307. — 
S. bakeri Symington, J. Mai. Br. R. As. Soc. 14 
(1936) 356, t. xx. — Fig. 7. 

Shrub or small to medium-sized tree. Twigs 
often stout, glabrous. Leaves glabrous, mostly 
faintly acuminate with cuneate or rounded base 
and undulate to crenate margin, elliptic, 5-15 by 
2-2 3 / 4 cm; nerves 9-14 pairs, meeting in an intra- 
marginal vein 2-4 mm from the margin ; petiole 
stout, 3-10 mm. Flowers in a subterminal, rarely 
terminal, raceme or panicle of racemes; axis 
pubescent to glabrous. Bracts and bracteoles 
glabrous, soon caducous, c. 8 and c. 5 mm long 
respectively. Pedicel at most 3 mm. Calyx glabrous, 
3-5 mm, sometimes becoming symmetric by 
tearing; lobes 2-3 mm, becoming longer by tearing 
apart. Corolla 8-10 mm. Stamens c. 100 or more. 
Disk 5-glandular, included the conical style base 
glabrous or soft hairy. Ovary glabrous, 1 1 / 2 -2 mm 
high; style glabrous, c. 5 mm. Fruit ellipsoid, c. 15 
by 8 mm ; stone smooth or with faint ridges, 
1-celled. Seeds not seen, but probably with straight 
embryo. 

Distr. Malesia: Malay Peninsula (Pahang: G. 
Tahan; Kuantan: G. Tapis), two collections. 

Ecol. Montane rain-forest, 1400-1650 m. Fl. 
June. 

Note. Closely allied to S. nivea, see there. 

47. Symplocos robinsonii Ridl. J. Fed. Mai. St. 
Mus. 8 (1917) 60; Noot. Leid. Bot. Ser. 1 (1975) 
276. 

Twigs tomentose, dark brown pubescent or 
(sparsely) appressedly pubescent or puberulous, 
glabrescent. Leaves sparsely long pubescent, 
appressedly fine dark-pilose or sparsely appres- 
sedly minutely pilose, glabrescent beneath, acute 
or acuminate with acute base and dentate, denticu- 
late or crenulate margin, narrowly to broadly 
elliptic, 3-9V2 by lV2-4cm; nerves 7-14 pairs. 
Flowers in an often branched raceme to 1 , 2 or 4 cm ; 
axis pubescent or appressedly puberulous. Bracts 
and bracteoles caducous, with same indument as 
axis, 1-2 and 3 / 4 to IV2 mm respectively. Pedicel to 
2 or 3(-4) mm long. Calyx pubescent to puberulous, 
often less hairy than ovary, 1-2 mm, the lobes 
V2-IV2 mm. Corolla 4-5 mm. Stamens 25-55. Disk 
with some hairs or shortly pilose, often the indu- 
ment hardly visible. Ovary with same indument as 
calyx or more hairy; style glabrous, or the base 
shortly pilose. Fruit ellipsoid, 7-10 by 3-6 mm; 
stone inconspicuously lengthwise grooved, 3-celled. 
Seeds 1-3, straight with straight embryo. 

Distr. Malesia: Sumatra. 



KEY TO THE VARIETIES 

1. Twigs tomentose a. var. robinsonii 

1 . Twigs not tomentose. 

2. Inflorescence to 4 cm long. Leaves 3-6 by 2-4 
cm (index IV4-2V4) . . . . b. var. latifolia 
2. Inflorescence 1-2 cm long. 
3. Twigs densely dark-brown pubescent. Leaves 
4-6V2 by lV 2 -372cm (index l 3 / 4 -2 3 / 4 ). 

c. var. pilosa 
3. Twigs sparsely appressed-pubescent or pube- 
rulous. Leaves 5-9'/2 by 2-3 cm (index 
2V4-3V2) d. var. angustifolia 

a. var. robinsonii. — Cf. Noot. Leid. Bot. Ser. 1 
(1975)277. — Fig. 7. 

Twigs tomentose. Leaves sparsely long-pubes- 
cent, especially on midrib and nerves beneath, ± 
elliptic, 4'/ 2 -9 by 2-3 3 / 4 cm; nerves 7-9 pairs; 
petiole 7-10 mm. Raceme to 2 cm, axis rusty 
pubescent. Bracts and bracteoles appressedly 
pubescent, Vj 2 and 1 mm long respectively. Pedicel 
to 3 mm. Calyx appressedly pubescent, c. 1 mm, 
the ± ovate lobes 3 / 4 -l mm long. Corolla c. 4 mm. 
Stamens 25-40. Disk with some hairs. Ovary ap- 
pressedly pubescent, c. l'^mm high; style with 
conical base, glabrous. Fruit 7 by 3 mm; stone 
3-celled. 

Distr. Malesia: Sumatra (Westcoast: G. 
Kerintji). 

Ecol. Gleichenia woodland in mountain forest, 
2200-2500 m. Fl. May, Aug. 

b. var. latifolia Noot. Leid. Bot. Ser. 1 (1975) 277. 
— Fig. 7. 

Treelet 6 m. Twigs (sparsely) appressedly pubes- 
cent, glabrescent. Leaves sparsely minutely appres- 
sedly pilose beneath, especially on midrib and 
nerves, or glabrous, shortly acuminate, 3-6 by 
2-4 cm ; nerves 7-8 pairs ; petiole 4-7 mm. Raceme 
branched, to 4 cm ; axis (sparsely) appressedly 
puberulous. Bracts and bracteoles ovate, 1-1 Va 
mm. Pedicel to 3(-4) mm. Calyx sparsely appres- 
sedly puberulous, lV 4 -2mm long, the ± semi- 
orbicular lobes 1-1 V4 mm long. Corolla c. 5 mm. 
Stamens 35-55. Disk with the conical style base 
pilose. Ovary with same indument as calyx, c. l'/ 4 
mm high; style glabrous, c. 4 mm. Fruit c. 10 by 
6 mm, blue-black. Seeds 1-3. 

Distr. Malesia: northern half of Sumatra (Gajo 
Lands: G. Losir; Westcoast: G. Kerintji). 

Ecol. In dense ericoid shrub-forest, 2000-3000 
(-3400) m. Fl. May-Aug. Flowers scentless. 

c. var. pilosa Noot. Leid. Bot. Ser. 1 (1975) 278. 
Twigs densely dark brown pubescent. Leaves 

appressedly fine dark pilose beneath, especially on 
midrib and nerves, acute to acuminate, ± elliptic, 
4-6V2 by lV2-3V2cm; nerves 7-10 pairs; petiole 
5-7 mm. Raceme to 1 cm ; axis appressedly brown 
pubescent. Bracts and bracteoles with same 
indument, IV2-2 and 1-1 '/ 2 mm respectively. 
Pedicel to 2 mm. Calyx sparsely fine puberulous, 
c. IV2 mm, the lobes semi-elliptic to ovate, c. 1 mm 
long. Corolla 4-5 mm. Stamens 30-45. Disk 
minutely pilose, hairs sometimes very inconspi- 
cuous. Ovary appressedly fine puberulous, 1V 4 - 
P/2 mm high; style glabrous, 4-5 mm. 



1977] 



Symplocaceae (Nooteboom) 



271 



Distr. Malesia: Sumatra (Westcoast: G. Merapi 
and G. Singalang). 

Ecol. Subalpine mountain forest, between lava 
boulders, 2500-2800 m. Fl. May-June. 

d. var. angustifolia Noot. Leid. Bot. Ser. 1 (1975) 
278. 

Twigs sparsely appressedly pubescent or puberu- 
lous. Leaves sparsely appressedly minutely pilose 
beneath, acuminate, 5-9 1 ! 2 by 2-3 cm ; nerves 9-14 
pairs; petiole 5—12 mm. Raceme to 2 cm; axis 
minutely appressedly puberulous. Bracts and 
bracteoles with same indument, ovate, c. 1 and 
3 ,' 4 mm respectively. Pedicel to 3 mm. Calyx less 
hairy than ovary, c. 1 mm long, the lobes ovate, 
1 2 - 3 4 mm. Corolla c. 4 mm. Stamens c. 35. Disk 
minutely pilose. Style glabrous. 

Distr. Malesia: Sumatra (Westcoast: G. Ophir 
= G. Talakmau). 

Ecol. Subalpine mountain forest, 1900-2700 m. 
Fl. May. 

48. Symplocos rubiginosa Wall. (Cat. 1831, 
n. Mil, nomen) ex DC. Prod. 8 (1844) 257; Miq. 
Fl. Ind. Bat. 1, 2 (1859) 466; Clarke, Fl. Br. Ind. 3 
(1882) 580; Brand, Pfl. R. Heft 6 (1901) 53; 
K. & G. J. As. Soc. Beng. 74, ii (1906) 247; Ridl. 
Fl. Mai. Pen. 2 (1923) 306; Noot. Leid. Bot. Ser. 1 
(1975) 279. — Lodhra rubiginosa Miers, J. Linn. 
Soc. Bot. 17 (1879) 299. — Fig. 7. 

Shrub, or tree to 30 m high and 50 cm 0. Twigs 
tomentose, pubescent, tomentellous or glabrous, 
rather thick (3-5 mm). Leaves sparsely appressedly 
pilose to more or less densely patently soft-villous 
beneath, especially on midrib and nerves, usually 
abruptly acuminate with cuneate base and finely to 
rather coarsely dentate margin, narrowly elliptic 
to obovate, 15-45 by 5 3 4-17cm; nerves 12-17 
pairs; petiole thickened, 10-25 mm. Flowers in a 
spike from wood beneath or between the leaves; 
in bud the inflorescence has the appearance of a 
short cone; axis pubescent to tomentellous, 
l-5(-8)cm. Bracts and bracteoles caducous as 
soon as the flower matures, ovate, boat-shaped, 
densely silky-pubescent, 3-5 and 2-3 mm respec- 
tively. Calyx appressedly puberulous to silky 
pubescent, often symmetrically torn, l'/ r 3mm, 
the lobes ' 2 -2 mm. Corolla sparsely (minutely) stiff 
hairy towards the outer base, 4-5 mm. Stamens 
60-100. Disk glabrous or sparsely pilose. Ovary 
pubescent to tomentellous or with same indument 
as calyx, 1-2 mm high; style glabrous or pilose, 
sometimes with thick conical pilose base. Fruit 
blue in vivo, ovoid to ellipsoid, sparsely short pilose 
to glabrous, 8-10 by 5 8 mm; stone lengthwise 
grooved, at one side with a deep transverse con- 
striction at ' 4 from the base. Seed I, once or twice 
and then S-shaped curved due to the constriction 
of the stone. 

Mali -in Sumatra, Malay Peninsula, and 
Borneo (rare in Kalimantan). 

Ecol. Moth in the lowland and in the hills, from 
sca-lcvcl to IKOOm, in primary and SCCOndar) 
mixed rain-forest, noi rarely in Diptcrocarp forest, 
along streamsidcs, on kcranga-., in luiiuni 

(once 

April), fr. J.i! I > I ruit remain white lor a long 
time, then turn through red to light blue when ripe 



Uses. The wood is very hard and used for house- 
building (Burk. Diet. 1935, 2115). 

Vern. Sumatra: lempaong kantjil, Palemb.; 
Malaya: pemasa, Sakai lang. ; Borneo: smuak, 
Sarawak, Land-Dayak. 

49. Symplocos salicioides Noot. Leid. Bot. Ser. 1 
(1975)280. 

Shrub 2 m, with pubescent twigs. Leaves faintly 
acuminate to sharply acute, with cuneate to round- 
ed base, pubescent beneath, narrowly elliptic, 
3V 2 -7 by s /«-lVaCm; nerves 6-8 pairs, rather 
inconspicuous, meeting in an intramarginal looped 
vein; petiole 3-4 mm. Spike 1 -flowered. Bracts and 
bracteoles pubescent, 2 and 1 mm long respectively. 
Calyx densely pubescent, divided into 1-1 ' 4 mm 
long triangular lobes. Corolla 2-2'/ 2 mm. Stamens 
15-20. Disk with the conical style base softly long- 
hairy. Ovary with same indument as calyx, 1 '/ 4 mm 
high ; style hairy for its lower half, c. 2 mm long. 
Fruit long ellipsoid, pubescent, 13 by 5 mm, only 
seen immature. 

Distr. Malesia: East New Guinea (Sepik area, 
once). 

Ecol. Lowland rain- forest, 1000 m. 

50. Symplocos sumatrana Brand, Pfl. R. Heft 6 
(1901) 62; Noot. Leid. Bot. Ser. 1 (1975) 283. — 
Fig. 7. 

Treelet 3 m. Twigs densely patently red-brown 
long-hairy or tomentose. Leaves softly pilose 
beneath, acuminate with rounded base and denticu- 
late margin, narrowly elliptic to ovate, 6-14 by 
2-4 cm; nerves 7-15 pairs, meeting in a looped 
much prominent intramarginal vein; petiole 
5-7 mm. Flowers in a spike or raceme of 2-4 cm; 
axis brown tomentose or spreadingly hairy. Bracts 
and bracteoles soon caducous, the first not seen, the 
latter appressedly long-hairy, c. 2" 2 mm long. 
Calyx divided into 5 appressedly pilose semi- 
elliptic 2 mm long lobes. Corolla c. 5 mm. Stamens 
45-70. Disk pulvinate, pilose. Ovary sericeous, 
c. 1 ' 2 mm high ; style with some hairs in the lower 
half, 2-5 mm. Fruit el